I r HARVARD UNIVERSITY Library of the Museum of Comparative Zoology The Great Basin Naturalist VOLUME 42, 1982 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University TABLE OF CONTENTS Volume 42 Number 1 - March 31, 1982 Utah flora: Rosaceae. Stanley L. Welsh 1 Seasonal foods of coyotes in southeastern Idaho: a multivariate analysis. James G. Mac- Cracken and Richard M. Hansen 45 Structure of Alpine plant communities near King's Peak, Uinta Mountains, Utah. George M. Briggs and James A. MacMahon 50 Observations on the reproduction and embryology of the Lahontan tui chub, Gila bi- color, in Walker Lake, Nevada. James J. Cooper 60 The prevalence of Echinococcus granulosus and other taeniid cestodes in sheep dogs of central Utah. Lauritz A. Jensen, Ferron L. Andersen, and Peter M. Schantz 65 Growth of juvenile American lobsters in semiopen and closed culture systems using for- mulated diets. S. R. Wadley, R. A. Heckmann, R. C. Infanger, and R. W. Mickel- sen 67 Diameter-weight relationships for juniper from wet and dry sites. T. Weaver and R. Lund 73 A description of Timpie Springs, Utah, with a preliminary survey of the aquatic macro- biota. Thomas M. Baugh, Michael A. Nelson, and Floyd Simpson 77 Vegetation and soil factors in relation to slope position on foothill knolls in the Uinta Basin of Utah. Miles O. Moretti and Jack D. Brotherson 81 Weather conditions in early summer and their effects on September blue grouse {Den- dragapus obscurus) harvest. Joy D. Cedarleaf, S. Dick Worthen, and Jack D. Brotherson 91 Description of the female of Phalacwpsxjlla hamata (Siphonaptera: Hystrichopsyllidae). R. B. Eads and G. O. Maupin ". 96 First record of pygmy rabbits {Brachijlagus idahoensis) in Wyoming. Thomas M. Camp- bell III, Tim W. Clark, and Craig R. Groves 100 Paspalum distichum L. var. indutum Shinners (Poaceae). Kelly Wayne AUred 101 Local floras of the Southwest, 1920-1980: an annotated bibliography. Janice E. Bowers . 105 The relation between species numbers and island characteristics for habitat islands in a volcanic landscape. Steven H. Carter-Lovejoy 113 Taxonomic studies of dwarf mistletoes {Arceuthobium spp.) parasitizing Pinus strobi- fomiis. Robert J. Mathiasen 120 Number 2 - June 30, 1982 Utah plant types— historical perspective 1840 to 1981— annotated list, and bibliography. Stanley L. Welsh 129 A new species of Cryptantha (Boraginaceae) from Nevada. Kaye H. Thome and Larry C. Higgins 196 New taxa of thistles {Cirsium: Asteraceae) in Utah. Stanley L. Welsh 199 A species of Cryptantha (Boraginaceae) dedicated to the memory of F. Creutzfeldt. Stanley L. Welsh 203 Algal populations in Bottle Hollow Reservoir, Duchesne County, Utah. Jeffrey Johansen, Samuel R. Rushforth, and Irena Kaczmarska 205 Herpetological notes from the Nevada Test Site. Wilmer w. Tanner 219 New species of American bark beetles (Coleoptera: Scolytidae). Stephen L. Wood 223 Temperature-related behavior of some migrant birds in the desert. George T. Austin and J. Scott Miller 232 Description of a new Phalacropsylla and notes on P. alios (Siphonaptera: Hys- trichopsyllidae). R. B. Eads and E. G. Campos 241 Vegetation of the mima mounds of Kalsow Prairie. Jack D. Brotherson 246 Occurrence and effect of Chrysomyxa pirolata cone rest on Picea pungens in Utah. Da- vid L. Nelson and Richard G. Krebil 262 Number 3 - September 30, 1982 Buccal floor of reptiles, a summary. Wilmer W. Tanner and David F. Avery 273 Western diamondback rattlesnake in southern Nevada: a correction and comments. Frederick H. Emmerson 350 Prevalence of Elaeophora schneideri and Onchocerca cervipedis in mule deer from cen- tral Utah. Lauritz A. Jensen, Jordan C. Pederson, and Ferron L. Andersen 351 Ecomorphology and habitat utilization of Echinocereus engebnannii and E. triglochi- diatiis (Cactaceae) in southeastern California. Richard I. Yeaton 353 First specimen of the spotted bat {Eudenna maciilatum) from Colorado. Robert B. Fin- ley, Jr., and James Creasy 360 Status of introduced fishes in certain spring systems in southern Nevada. Walter R. Courtenay, Jr., and James E. Deacon 361 A new species of Penstemon (Scrophulariaceae) from the Uinta Basin of Utah and Colo- rado. John Larry England 367 Phalacropsis dispar (Coleoptera: Phalacridae), an element in the natural control of na- tive pine stem rust fungi in the western United States. David L. Nelson 369 Species-habitat relationships in an Oregon cold desert lizard community. David F. Werschkul 380 Prehminary index of authors of Utah plant names. L. Matthew Chatterley, Blaine T. Welsh, and Stanley L. Welsh 385 Nest site selection in raptor commimities of the eastern Great Basin desert. Dwight G. Smith and Joseph R. Murphy 395 Invertebrate faunas and zoogeographic significance of lava tube caves of Arizona and New Mexico. Stewart B. Peck 405 Additions to the vascular flora of Montana and Wyoming. Robert W. Lichvar and Rob- ert D. Dom 413 Number 4 - December 31, 1982 Ants of Utah. Dorald M. Allred 415 Vegetal responses and big game values after thinning regenerating lodgepole pine. D. D. Austin and Philip J. Urness 512 Habitat manipulation for reestablishment of Utah prairie dogs in Capitol Reef National Park. Rodney L. Player and Philip J. Urness 517 Effects of defoliation on reproduction of a toxic range plant, Zigadenus paniculatus. V. J. Tepedino 524 Distribution and relative abundance of fish in Ruth Reservoir, California, in relation to environmental variables. Steven Vigg and Thomas J. Hassler 529 Temperature and salinity relationships of the Nevadan rehct dace. Steven Vigg 541 Observations on woundfin spawning and growth in an outdoor experimental stream. Paul Greger and James E. Deacon 549 Early development of the razorback sucker, Xyrauchen texanus (Abbott). W. L. Minck- ley and Eric S. Gustafson 553 Behavior and habitat preferences of ring-necked pheasants during late winter in central Utah. Jeffrey G. Skousen and Jack D. Brotherson 562 Rhythm of fecal production and protein content for black-tailed jackrabbits. William D. Steigers, Jr., Jerran T. Flinders, and Susan M. White 567 Prairie dog colony attributes and associated vertebrate species. Tim W. Clark, Thomas M. Campbell III, David G. Socha, and Denise E. Casey 572 Distribution of the moss family Grimmiaceae in Nevada. Matt Lavin 583 Insular biogeography of mammals in the Great Salt Lake. Michael A. Bowers 589 Dorsal hair length and coat color in Abert's squirrel {Sciurus aherti). Denis C. Hancock, Jr., and Donald J. Nash 597 The raccoon, Procyon lotor, in Wyoming. E. Blake Hart 599 Intercanine crown distances in red foxes and badgers. E. Blake Hart 601 FHE GREAT BASIN NATURALIST Volume 42 No. 1 March 31,1982 Brigham Young University MUS. ' 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, Id- aho 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. Scliolarly 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. 6-82 650 59839 ISSN 017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 42 March 31, 1982 No. 1 UTAH FLORA: ROSACEAE Stanley L. Welsh' Abstract.- A revision of the rose family, Rosaceae, is presented for the state of Utah. Included are 115 species and 9 varieties of indigenous and introduced plants in 35 genera. A key to the genera and species is provided, along with detailed descriptions, distributional data, and pertinent comments. Proposed as a new taxon is Crataegus doug- lasii Lindl. var. duchesnensis Welsh. New combinations include Potentilla concinna Richards var. bicrenata (Rydb.) Welsh & Johnston; P. concinna var. modesta (Rydb.) Welsh & Johnston; P. concinna var. proxima (Rydb.) Welsh & Johnston; P. glandulosa Lindl. var. micropetala (Rydb.) Welsh & Johnston; P. ovina J. M. Macoun var. decurrens (Wats.) Welsh & Johnston; P. pensylvanica L. var. paticijuga (Rydb.) Welsh & Johnston. This paper is one of a series of taxonomic revisions leading to a definitive treatment of the flora of Utah. The rose family is of mod- erate size in the state, but it is important for the indigenous species that comprise portions of the plant communities of substance. The family is important, too, for its introduced or- namental and fruit plants. Apples, pears, raspberries and relatives, cherries, peaches, plums, apricots and relatives, and straw- berries are important products of orchards and gardens. Introduced taxa number 44, or 38 percent of the total rosaceous flora reported herein. Half of those taxa belong to genera not rep- resented in the indigenous flora. Practically all of the introduced taxa are cultivated orna- mental or fruit plants. It is a remarkable fam- ily to have so few weedy species. Only Geum and Potentilla support species v/hich are weedy, but some of the cultivated fruit plants or their rootstocks that escape can become problems. The Himalayan blackberry escapes and persists as a spiny bramble in lower ele- vation agricultural regions. The most difficult and largest genus in the family in Utah is Potentilla. That genus pre- sents an amazing array of intergrading taxa, which have been subjected to a series of in- terpretations. The extreme interpretations in- volve recognition of an infinite number of taxa at species rank on the one hand, and the subjugation of most of these in synonymy of broadly defined species on the other. The treatment of Potentilla presented here is the result of collaboration between me and Barry C. Johnston of the U.S. Forest Service, in Denver. It strikes a position somewhere be- tween the extremes, and represents a com- promise between the views of the authors. We have chosen to avoid the use of the seg- regate generic names, except to indicate their position in the key to the species, and to present the synonymy that will allow use by those who might be so inclined. 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 2 Great Basin Naturalist Vol. 42, No. 1 RosACEAE some), commonly showy; stamens 5 to nu- merous; pistils 1 to many, of 1 carpel, or of 5 Rose Family connate or distinct carpels enclosed in the hypanthium; fruit an achene, follicle, drupe, Annual, biennial, or perennial herbs, pome, aggregate, hip, or accessory. The rose shrubs, or trees; leaves alternate or basal (and family is both large and complex. The diver- still alternate) or less commonly opposite, sity of fruit type reflects the many morpholo- simple or pinnately to palmately compound, gical differences in structure of the gynoe- mostly deciduous, stipulate or rarely exstipu- cium in this assemblage. Suggestions by some late; flowers perfect or imperfect, regular, workers that the group should be segregated complete or incomplete, perigynous to epi- into more than one family is not without gynous, borne singly or in racemose, corym- merit. They are held together by the pres- bose, umbellate or cymose clusters; sepals ence of the hypanthium on which the per- usually 5 (more in some), often bearing brae- ianth and stamens are displayed. This is a teoles alternate with the lobes, borne with complex structure, with several possible ori- petals and stamens on margin of a hypan- gins, and might fail ultimately as a diagnostic thium; petals usually 5 (lacking or more in character. 1. Plants annual, biennial, or perennial herbs Subkey I - Plants trees, shrubs, or subshrubs Subkey II Subkey I. 1. Petals lacking; flowers numerous, borne in dense spikes; leaves pinnately ^°"^P«^^ Sanguisorba - Petals present; flowers not both numerous and borne in spikes 2 2(1). Leaves bi- or triternately dissected into linear segments; petals white- plants of Piute, Beaver, and Sevier counties Chamaerhodos - Leaves various, but not bi- or triternately dissected into linear segments; petals white, yellow, or pink ' o 3(2). Flowers solitary on scapose peduncles; leaves simple, crenate; fruit of plumose achenes; sepals and petals 8-10 each Druas - Flowers usually more than 1; leaves compound or lobed, rarely simple; fruit not of plumose achenes; sepals and petals usually 5 each ' 4 4(3). Bractlets lacking between the sepals; flowers with a stalked receptacle; hypantiiium funnelform; plants of Washington Co Purpusia - Bractlets present, alternating with the sepals; flowers with a sessile receptacle; hypanthium not funnelform ' 5 5(4). Leaflets tridentate apically, entire along the sides; stamens 5; plants prostrate or mat forming, of high elevations Sibbaldia - Leaflets variously tooflied or lobed, but not regularly tridentate apically; stamens 5, 10, or more; plants of various habit and habitat ' 6 6(5). Leaves trifoliate; plants with well-developed stolons; flowers white; receptacle ripening into an accessory fruit .'. Framria - Leaves mostly with more than 3 leaflets, but, if trifoliolate, then lacking stolons; flowers mostly yellow; receptacle not ripening 7 7(6). Leaflets very numerous, mostly less than 6 mm long; petals usually clawed Ivesia - Leaflets 3-15 (rarely more), commonly much more than 6 mm long; petals sessile o March 1982 Welsh: Utah Flora: Rosaceae 3 8(7). Leaves palmately or pinnately lobed or compound, not lyrate pinnatifid; styles at maturity not elongate and conspicuous Potentilla — Leaves pinnately lobed or compound or more usually lyrate-pinnatifid; styles at maturity elongate and conspicuous Geum Subkey IL 1. Leaves compound 2 — Leaves simple 7' 2(1). Stems and/or leaves armed with prickles or spines 3 — Stems and leaves lacking prickles or spines 4 3(2). Pistils several, enclosed within a fleshy hypanthium; fruit a hip; petals very showy Rosa — Pistils several to many, on an elongate receptacle; fruit an aggregate; petals not especially showy Rubiis 4(2). Leaves bipinnately compound, the ultimate segments 0.5-1.5 mm long; herbage glandular-stellate, aromatic Chamaebatiaria — Leaves once pinnately compound, the leaflets much longer than 1.5 mm; herbage not glandular-stellate 5 5(4). Leaflets 3-7; leaves 1.5-3.5 cm long; flowers yellow; low shrub Potentilla — Leaflets 7-15 or more; leaves 5-20 cm long or more; flowers white to cream; moderate shrubs to small trees 6 6(5). Ovary superior; stamens 20 or more; leaflets 13-23; cultivated shrubs Sorbaria — Ovary inferior; stamens 15-20; leaflets 9-15; indigenous shrubs or cultivated trees Sorbus 7(1). Leaves opposite; petals lacking; intricately branched, low desert shrubs of southern and southeastern Utah Coleogyne — Leaves alternate; petals present (lacking in Cercocarpus); plants of various habits and habitats 8 8(7). Shrubs low, mat forming; flowers solitary or in dense spikes on leafless or merely bracteate scapes • " — Shrubs or small trees, never mat forming; flowers various, but neither scapose nor subscapose 1^ 9(8). Flowers solitary, the sepals and petals mostly 8-10 each; leaves crenate; plants of alpine tundra Dryas — Flowers in dense spikes, the sepals and petals commonly 5 each; leaves entire; plants of rock surfaces at low to moderate elevations Petrophytum 10(8). Pistils superior, the 1 to several separate or partially connate; ovaries not adnate to the hypanthium; fruit a drupe, aggregate, achene, follicle, or capsule .... 11 — Pistils inferior, the 3- to 5-carpellate ovaries adnate to the hypanthium; fruit a 21 pome ^^ 11(10). Flowers inconspicuous; petals lacking; leaves entire and evergreen (except in C. lywntanus) Cercocarpus — Flowers showy, though small in some; petals present; leaves mainly toothed or lobed, often deciduous 12 4 Great Basin Naturalist Vol. 42, No. 1 12(11). Pistil 1; fruit a drupe; leaves commonly with glands at base of blade or on petiole p,^„j^^ — Pistils 1 to many; fruit not a drupe; leaves not gland-bearing 13 13(12). Leaves pinnately veined, the lobes, if any, pinnate 14 — Leaves palmately veined, the lobes palmately arranged or flabellate 16 14(13). Flowers yellow, sohtary, terminating branches of the current year Kerria — Flowers white to pink or lavender, borne in corymbs, panicles or racemes 15 15(14). Flowers borne in racemes, 1.5 cm wide or more; petals 6-12 mm long Exochorda — Flowers borne in corymbs or panicles, less than 1 cm wide; petals 6-15 mm lo"g Spiraea 16(13). Flowers large, 2 cm broad or more, in few-flowered cymes; fruit an aggregate . Rubus — Flowers commonly less than 2 cm broad, solitary or in corymbs or panicles 17 17(16). Flowers numerous, borne in panicles Holodiscus — Flowers borne singly or in few- to many-flowered corymbs 18 18(17). Flowers borne in umbellate corymbs; leaves broad and thin, commonly 1-6 cm wide or more Physocarpus — Flowers borne singly or in corymbose racemes; leaves thickish, seldom to 1 cm wide |Q 19(18). Pistils numerous; petals white; leaf lobes tightly revolute; plants of low elevations in southern Utah Fallupia — Pistils 1-5 (rarely more); petals white to cream or pale yellowish; leaf lobes not tightly revolute; plants of broad distribution 20 20(19). Pistils 1 or 2; styles not plumose; leaves usually 3-lobed Purshia — Pistils commonly 5 (or more); styles plumose at maturity; leaves commonly 5-to7-lobed Cowania 21(10). Stems armed with thorns or spines 22 — Stems unarmed 24 22(21). Leaves evergreen, crenate-serrate; pomes commonly orange; petals white, less than 4 mm long Pyracantha — Leaves deciduous, serrate or doubly serrate; pomes variously colored, rarely orange; petals more than 5 mm long 23 23(22). Shrubs to 2 m tall (generally less); flowers 20-45 mm broad; fruit over 2 cm ek Chaenomeles — Shrubs or small trees to 5 m tall or more; flowers 9-18 mm broad; fruit less than 1.5 cm thick '. Crataegus 24(21). Leaves entire or essentially so 25 — Leaves serrate to doubly serrate (see also Peraphyllum) 27 25(24). Leaves ovate to cordate ovate, 1.5-5 cm wide or more; pomes clothed with a villous tomentum Cydonia — Leaves variously shaped, less than 1.5 cm wide; pomes glabrous 26 26(25). Shrubs to 1.5 m tall or more, indigenous; leaves narrowly elliptic; fruit an acrid PO"^e Peraphyllum — Shrubs of various height, cultivated; leaves ovate to obovate or oblanceolate; pomes mealy, nonacrid Cotoneaster March 1982 Welsh: Utah Flora, Rosaceae 27(24). Flowers white, in racemes; plants indigenous, rarely cultivated; leaves prominently toothed toward the apex Amelanchier — Flowers white or otherwise, in corymbs; plants cultivated, sometimes escaping; leaves toothed or lobed throughout 28 28(27). Leaves deeply or at least prominently lobed Sorbus — Leaves moderately, if at all, lobed 29 29(28). Shrubs to 2 m tall (generally less); flower solitary or sessile in corymbose clusters...;. Chaenomeles — Shrubs or trees to 7 m tall or more; flowers pedicellate in corymbose or umbellate clusters ^^ 30(29). Flowers in umbels; styles connate at the base; fruit with few if any stone cells, apple-shaped ^«^"« — Flowers in corymbs; styles free; fruit with stone cells, mostly pear shaped Pyrus Amelanchier Medic. sepals 5, persistent; petals 5, white; stamens usually 10 or more; pistil 1, the ovary inferi- Shrubs or small trees with unarmed or, usually 5-loculed (appearing as 10); styles branches; leaves alternate, simple, not lobed; 2-5, the stigmas capitate; fruit a reddish to stipules linear, caducous; flowers perfect, purplish, often glaucous, pome, regular, borne in racemes; hypanthium short, Jones, G. N. 1946. American species of Ame- with a glandular disk on the inner surface; lanchier. 111. Biol. Monogr. 20: 1-126. 1 Leaves mainly over 2.5 cm long; petals mostly 9-15 mm long; style commonly 5 A. alnifolia — Leaves mainly less than 2.5 cm long; petals 5-10 mm long; styles 2-4 (rarely 5) A. utahensis Amelanchier alnifolia (Nutt.) Nutt. Ser- viceberry, Shadbush, Saskatoon. [Aronia alni- folia Nutt.; Pynis alnifolia (Nutt.) Lindl.; A. canadensis var. alnifolia (Nutt.) T. & G.; A. canadensis var. pumila Nutt. in T. & G.; A. pumila (Nutt.) Roem.; A. alnifolia var. cu- sickii (Fern.) C. L. Hitchc; A. polycarpa Greene]. Low shrubs to small trees, mostly 2-5 m tall; leaves petiolate, mainly 20-50 mm long, 15-40 mm broad, oval to oblong, acute to rounded or subcordate basally, roimded to truncate apically, serrate near the apex, glabrous or hairy on one or both sides; flowers in short racemes; sepals 2.8-4.6 mm long; petals 9-15 mm long, 3.3-5.8 mm wide, spatulate-oblanceolate, white to pinkish; styles 5 (or 4); fruit purplish to black purple, glaucous, subglobose, 6-14 mm long, pala- table. Streamsides, meadows, and mountain slopes at 1500 to 2900 m in sagebrush, moun- tain brush, aspen and mixed conifer woods in Box Elder, Cache, Davis, Garfield, Iron, Mil- lard, Morgan, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, Tooele, Uintah, Utah, Weber, and Washington counties; Alaska and Yukon east to Hudson Bay and south to California, Arizona, New Mexico, and Nebraska. Attempts to segregate the var- ious proposed infraspecific taxa among our Utah materials are fraught with difficulties not easily overcome, even by application of mechanical and arbitrary keys. Pubescence or its absence and the position of that pu- bescence form the basis of the main segre- gates. The feature of pubescence seems to be so variable, not only within A. alnifolia, but within A. utahensis (q.v.), that it might in- dicate a response to ecological conditions rather than genetic affinities. More work is indicated; 66 (iii). Amelanchier utahensis Koehne Utah Ser- viceberry. [A. hakeri Greene; A. oreophila A. Nels.; A. utahensis ssp. oreophila (A. Nels.) Clokey; A. florida var. oreophila (A. Nels.) R. J. Davis; A. crenata Greene; A. elliptica A. Nels.; A. prunophil'a Greene; A. rubescens Greene; A. utahensis var. cinerea Goodding, type from Washington Co.]. Low to large 6 Great Basin Naturalist Vol. 42, No. 1 shrubs, mostly 0.5-4 m tall, intricately available, the best of characteristics fail sin- branched, often in dense clumps; leaves pe- gly and often in combination as well. Because tiolate, mainly 10-27 mm long, 6-27 mm of the trends indicated by leaf and petal size wide, oval to ovate, oblong, or elliptic, acute and other features, it seems best to treat A to rounded or subcordate basally, rounded to utahensis apart from the tangled morphology truncate or less commonly acute apically, of A. a/nt/o/fa. Additionally, variation within serrate near the apex, hairy on one or both die utahensis assemblage is as great as (or sides rarely glabrous; sepals 1-3 mm long; greater) than that known to occur in the alni- petals 5.2-10 mm long, 1.8-4.2 mm wide, folia materials; 193 (xxvi). spatulate-oblanceolate to elliptic, white, cream or pinkish; styles 2-4 (5); fruit purplish Cercocarpus H. B. K. or pinkish, 5-12 mm long, palatable or dry Shrubs or small trees with unarmed and hardly edible. Streamsides, dry slopes, or branches and very dense wood; leaves alter- thickets m sagebrush, grassland, mountain nate, simple, entire or toothed; stipules small mahogany mountain brush, pinyon-juniper, adnate to petiole; flowers perfect regular' X?' 021^°""^^'^ P'"^ communities at borne solitary or in small clusters, terminal or 900 to 2800 m in all counties in Utah (type axillary; hypanthium trumpetlike, with a de- from Leeds, Washington Co.); Washington to ciduous apical portion; sepals 5; petals lack- Montana and south to Baja California, Ari- ing; stamens 10 or more, borne in 2 or 3 zona, New Mexico, and Texas. Segregation of [Bsw; pistils 1, of 1 carpel; style terminal- all specimens in the alnifolia-utahensis com- fruit an achene, with the elongate plumose piex IS difficult if not impossible. Diagnostic style persisting. features show overlap, and, although trends Martin, F. L. 1950. A revision of Cerco- are apparent in the vast amount of material carpus. Brittonia 7: 91-111. 1. Leaves deciduous, toothed, not especially re volute C. montanus - Leaves evergreen, entire or toothed, decidedly revolute 2 2(1). Leaves (at least some) toothed; a hybrid C. montanus X C. ledifolius — Leaves entire „ 3(2). Leaves elliptic, commonly 12-30 mm long or more; shrubs or small trees mostly of middle and higher elevations C ledifolius Leaves linear to narrowly oblong, usually less than 12 mm long; low intricately branched shrubs of lower middle and lower elevations C. intricatus Cercocarpus intricatus Wats. Dwarf Nevada, California, and Arizona. Pubescence Moimtain Mahogany. [C. ledifolius var. in- of leaves varies in form from strigose-pilose tncatus (Wats.) Jones; C. intricatus var. vil- to crinkly-hairy, or is lacking. Plants with pi- losus Schneid., type from Deep Creek, Juab lose leaves form the basis of var. pilosa, but (?) Co.; C. arizonicus Jones]. Shrubs mostly the feature does not seem to be correlated 0.5-2 m tall, intricately branched; leaves with any other. Leaves are heavily cutinized 3-18 mm long, 0.8-1.4 mm wide, oblong to in plants from Kane and Washington coun- linear (rarely elliptic), tightly revolute, ties; 53(xiii). glabrous, strigose-pilose, or villous, co- Cercocarpus ledifolius Nutt. in T. & G. riaceous and persistent; flowers 3.2-8.7 mm Curl-leaf Mountain Mahogany. [C. ledifolius long; sepals 0.6-1.2 mm long; stamens 10-20; var. intercedens Schneid.; C. ledifolius var. tails of achenes 1-3 cm long. Rimrock, cliffs, intercedens f. subglaher Schneid., type from and slopes in desert shrub, pinyon-juniper Slate Canyon, Utah Co.; and f. hirsutus and mountain brush communities at 1370 to Schneid., type from Ogden, Utah]. Shrubs or 2400 m in Beaver, Emery, Garfield, Grand, small trees, mainly 2-5 m tall; leaves 10-42 Juab, Kane, Millard, San Juan, Sanpete, Se- mm long, 2-14 mm wide, elliptic to oblong, vier, Uintah, Utah (type from American Fork the margin only revolute, pubescent to Canyon), Washington, and Wayne counties; glabrous, coriaceous, persistent; flowers 7-10 March 1982 Welsh: Utah Flora, Rosaceae mm long; sepals 1.2-2.1 mm long; stamens 20-30; tails of achenes 4.5-8 cm long. Moun- tain brush, pinyon-juniper, aspen, and spmce- fir commimities, often in stands, at 1400 to 3000 m in Beaver, Box Elder, Cache, Carbon, Emery, Garfield, Grand, Iron, Juab, Millard, Morgan, Piute, Rich, Sanpete, Summit, Tooele, Uintah, Utah, Washington, Wasatch, and Weber counties; Washington to Montana and southward to California, Arizona, and Colorado. Hybrids are known between C. ledifolius and C. montanns. They are easily discerned by the coriaceous persistent tooth- ed leaves. They occur as scattered individuals in places of contact between the parental types. Similarly, putative hybrids involving C. ledifolius and C. intricatus are known. Longer, very narrow, and markedly revolute leaves mark those apparent hybrids; 89(xi). Cercocarpus montanus Raf. Alder-leaf Mountain Mahogany. [C. betuloides Nutt. in T. & G.; C. betidaefolius Nutt. ex Hook.; C. parviflorus var. glaber Wats.; C. parviflorus var. betuloides (Nutt.) Sarg.; C. montanus var. glaber (Wats.) Martin; C. parvifolius var. mini77ius Schneid.; C. flabellifolius Rydb., type from Glenwood, Sevier Co., Utah]. Shrubs, or less commonly, small trees com- monly 1.2-4 m tall; leaves short-petiolate, the blade obovate to oblanceolate or orbicu- lar, 6-44 mm long, 5-23 mm wide, crenate- serrate, glabrous above, pubescent beneath (sometimes glabrous), deciduous; flowers 9.5-17.5 mm long; sepals 0.9-1.7 mm long; stamens 25-40, the anthers hairy; tails of ach- enes 3-10 cm long. Mountain brush, sage- brush, grassland, pinyon-juniper, aspen, and mixed conifer communities at 1400 to 2800 m throughout Utah; Oregon to Wyoming and south to Mexico. This and other species of Cercocarpus are valuable browse plants for wildlife and domestic livestock. They are components of wild seed mixtures in rec- lamation attempts; 91(xiv). Chaenomeles Lindl. Shrubs, usually armed with thorns; leaves alternate, simple, serrate, stipules large, de- ciduous; flowers perfect, regular, solitary or 2-5 or more in sessile clusters; hypanthium short; sepals not persistent; petals 5 (some- times more), variously colored; stamens 20 or more; pistil 1, the ovary inferior, usually 5-lo- culed; styles 5, joined at the base; the stigmas capitate; fruit a pome of moderate size. Branchlets with vernicose small scars left by deciduous short hairs; flowers orange scarlet, about 2.5 cm wide; plants mostly less than 1 m tall C. japonica Branchlets smooth, lacking hair scars; flowers variously colored, over 2.5 cm wide; plants commonly more than 1 m tall C. speciosa Chaenomeles japonica (Thunb.) Lindl. ex Spach. Japanese Quince. [Pijrus japonica Thunb.; Cydonia japonica (Thunb.) Pers; Cij- donia lagenaria Lois.; Cydonia japonica var. lagenaria (Lois.) Makino; Chaenomeles lage- naria (Lois. Koidz.]. Shrubs to 1 m tall (rarely more); the short branchlets often modified as thorns, the young branchlets with deciduous short hairs leaving verrucose scars on falling; leaves obovate, 20-50 mm long, 8-35 cm wide, obtusely serrate, obtuse to subacute apically; flowers orange scarlet, about 2.5 cm wide; fmit subglobose, about 3 cm thick. Cultivated ornamental in Carbon, Salt Lake, and Utah coimties; introduced from Japan; 4(i). _ Chaenomeles speciosa (Sweet) Nakai Flowering Quince. [Cydonia speciosa Sweet; C. lagenaria Koidz. not (Lois.) Koidz.]. Shrubs to 2 m tall, the short branchlets often modified as spines, glabrous or with short de- ciduous hairs leaving no scars on falling; leaves oblong to ovate or lanceolate, 22-65 mm long, 12-35 mm wide, sharply serrate, acute to subacute apically; flowers scarlet to white or red, 2.5-3.5 cm wide; fruit sub- globose to pyriform, 2-5 cm thick. Culti- vated ornamental in Juab, Salt Lake, and Utah counties; introduced from China; 3(i). Chamaebatiaria (Porter) Maxim. Aromatic shrubs, unarmed; leaves alter- nate, bi- or tripinnately compound, the her- bage stellate-pubescent; stipules herbaceous, more or less persistent; flowers perfect, Great Basin Naturalist Vol. 42, No. 1 regular, showy, borne in terminal panicles; hypanthium turbinate; sepals 5, persistent; petals 5, white; stamens many; pistils 5, more or less connate below, the ovary superior; styles 5; fruit of follicles. Chamaebatiaria millefolium (Torr.) Max- im. Fern Bush, Desert Sweet. [Spiraea mille- folium Torr.]. Shrub 8-20 dm tall (rarely more), the stems and herbage glandular and stellate-pubescent when young; leaves 0.9-6.7 cm long, 0.4-1.8 cm wide, oblong to lanceolate in outline, with 8-24 pairs of pin- nae, these again pinnate, the tertiary seg- ments again pinnatifid; panicles 3-15 cm long; flowers 0.8-1.5 cm wide; sepals ovate to lanceolate, 3-5 mm long, green; petals white, 2.5-5 mm long and about as broad; follicles 4-6 mm long, few seeded. Sagebrush^ mountain brush, aspen, limber pine, and spruce-fir communities at 1800 to 2900 m in Beaver, Box Elder, Garfield, Iron, Juab, Kane, Millard, Piute, Tooele, and Washing- ton counties; Oregon, Idaho, Wyoming, and south to California and Arizona; 34 (vii). Chamaerhodos Bunge Plants biennial or short-lived perennial herbs; leaves alternate and in a basal rosette, bi- or triternately divided, the segments nar- row; stipules foliose, narrowly oblong, simple or divided, persistent; flowers perfect, regu- lar, borne in bracteate, corymbose cymes; hy- panthium cup shaped, long-hairy within; sep- als 5; petals 5; stamens 5, borne at the base of the petals; pistils 5 (rarely fewer), distinct, the ovaries superior, each 1-loculed; styles 1 per pistil, the stigma capitate; fruit an achene. Chamaerhodos erecta Bunge in Ledeb. American Chamaerhodos. [C. erecta var. nut- tallii T. & G.; C. erecta ssp. nuttallii (T. & G.) Hulten; C. nuttallii (T. & G.) Rydb.]. Plants erect, mostly 7-28 (30) cm tall, from a tap- root and a basal rosette, the stems freely branched above the base; leaves mostly 7-40 mm long, the ultimate segments linear to ob- long, sparingly long-hairy; cymes equaling 1/4 to 1/2 the plant height; flowers short-pedicel- late, inconspicuous; sepals 1.2-2.5 mm long, triangular, sparsely hirsute; petals white! equaling or slightly longer than the sepals; achenes 1.2-1.5 mm long, glabrous, grayish. Igneous gravel and sandy loam in sagebrush, grassland, and alpine tundra at 2745 to 3355 m in Piute, Sevier, and Wayne (?) counties; Alaska and Yukon east to Michigan and south to Colorado and North Dakota; Asia. Our materials are assignable to var. parviflora (Nutt.) C. L. Hitchc. [Sibbaldia erecta var. parviflora Nutt.); 4 (iv). CoLEOGYNE Torr. Shrubs, the stems intricately branched, spinescent; leaves opposite, fasciculate on spur branchlets, entire, coriaceous, persistent; flowers perfect, regular, solitary and terminal on spur branchlets, subtended by paired, tri- fid bracts; hypanthium cup shaped, co- riaceous, persistent; sepals 4, persistent; pet- als 0; stamens 20-40, basally inserted on outside of tubular sheath enclosing ovary; pistils 1, with a lateral, twisted, exserted, per- sistent style pubescent at base; fruit a glabrous achene. Coleogyne ramosissima Torr. Blackbrush. Rounded shrubs, 3-12 dm or more tall, with divaricate branches; leaves 3-12 mm long, mainly 0.8-1.5 mm wide, narrowly oblan- ceolate, obtusish and commonly mucronate apically, strigose with malpighian hairs; se- pals 4.5-6.5 (8) mm long, ovate to lanceolate, malpighian hairy and red brown dorsally, glabrous and yellowish ventrally; sheaths membranous, tapering to 5-toothed apex, silky-hairy within, glabrous without; achene ovate, curved, glabrous, 5-8 mm long. Shal- low sandy to clay soils in blackbrush and warm desert and shrub communities at 760 to 1830 m in Emery, Garfield, Grand, Kane, San Juan, and Washington counties; Nevada and Colorado south to California and Arizona- 35(v). COTONEASTER Mcdic. Shrubs, unarmed, erect to arcuate or hori- zontal, deciduous or evergreen; leaves alter- nate, simple, entire; stipules linear, de- ciduous; flowers perfect, regular, solitary or in cymes terminating lateral branches; hy- panthium short, persistent; sepals 5; petals 5, white or pink; stamens 10-20; pistil 1, the ovary inferior, 2- to 5-loculed, the styles 2-5; fruit a pome. Note: Members of this genus March 1982 Welsh: Utah Flora, Rosaceae 9 are cultivated widely in Utah as ornamentals. They have potential for use in reclamation and stabilization projects and will probably be maintained as a portion of our introduced flora. The genus has some 50 species dis- tributed in Eurasia, and many more are in cultivation than are treated herein. The spe- cies keyed below are merely representative. 1. Leaves mainly 5-12 mm long; flowers usually solitary 2 — Leaves mainly 15-100 mm long or more; flowers commonly several to many 3 2(1). Petals spreading, white; shrubs spreading; leaves evergreen C. microphylla Lindl. — Petals erect, pink; plants depressed-horizontal; leaves half evergreen C. horizontalis Decne. 3(1). Petals erect, obovate, pinkish or white; fruit red or black 4 — Petals spreading, suborbicular, white; fruit red 5 4(3). Fruit black; at least some leaves more than 2.5 cm long C. aciitifolia Turcz. — Fruit red; leaves less than 2.5 cm long C. dielsiana Pritz. 5(3). Leaves, glabrate at maturity C. multiflora Bunge — Leaves persistent, white to rusty tomentose beneath 6 6(5). Leaves white-tomentos ebeneath, commonly 1-3 cm long C. pannosa Franch. — Leaves rusty-tomentose beneath, or finally glabrate, often at least some over 3 cm long C salicifolia Franch. Note: Cotoneaster species are not described due to lack of adequate specimens in her- baria. Much work on cultivated plants is necessary. Cowania D. Don Shrubs or small trees; leaves alternate, pin- natifid, coriaceous, glandular-dotted; stipules minute, triangular, persistent; flowers per- fect, regular, solitary, terminal on spur branchlets; hypanthium funnelform, per- sistent; sepals 5; petals 5, white to cream or yellowish; stamens many; pistils 4-12 (com- monly 5), long-hairy, the style terminal, plu- mose, persistent and elongate in fruit; fruit an achene. Cowania mexicana D. Don Cliff-rose. Much branched shrubs or small trees, mainly 0.6-3.5 m tall with shreddy bark and glandu- lar branchlets; leaves 3-15 mm long, cu- neate-flabellate, mainly 5-lobed, glandular- punctate and green above, white-tomentose beneath; pedicels 2-8 mm long; sepals 4-6 mm long, ovate; petals 5-9 mm long, white to cream or yellowish; pistils commonly 5; styles plumose 2-6 cm long or more in fruit. Blackbrush, live oak, pinyon-juniper, pon- derosa pine, desert peach, mixed grass-desert shrub, and mountain brush communities at 975 to 2745 m in Beaver, Box Elder, Carbon, Emery, Garfield, Grand, Juab, Kane, Millard, Salt Lake, San Juan, Sevier, Tooele, Utah, Washington, and Wayne counties; Nevada, Colorado, Arizona, California, New Mexico, and Mexico. Our materials belong to var. stansburiana (Torr.) Jeps. [C. stansbiiriana Torr.]. This species forms intergeneric hy- brids with Purshia tridentata (q.v.). The prob- lem has been investigated by Stutz and Thomas (1964. Evolution 18:183-195); 112(xxiv). Crataegus L. Small, deciduous trees or shrubs, com- monly armed with thorns; leaves alternate, simple, serrate to doubly serrate, lobed in some; stipules small, adnate to the petiole, glandular-serrate, deciduous; flowers perfect, regular, in corymbose cymes terminating short lateral branchlets; hypanthium short, free above the ovary; sepals 5, tardily de- ciduous; petals 5, white or pink; stamens (5) 10-25 or more, the filaments filiform; pistil 1, the ovary inferior, 1- to 5-loculed, the styles 1-5; fruit a pome. Only one taxon is widespread in Utah, growing as a portion of the indigenous flora. The following key con- tains other native and widely cultivated or escaping taxa. Others are present in cultiva- tion but are excluded. 10 1. 2(1). 3(2). Great Basin Naturalist Vol. 42, No. 1 Leaves deeply 3- to 7-lobed; styles 1-3; fruits with 1 or 2 seeds; plants ornamental, cultivated and escaping C. monopuna Leaves serrate to doubly serrate or somewhat lobed; styles 2-5; plants indigenous 2 Leaves mostly more than twice as long as broad; plants common and widespread; fruit black C. douglasii Leaves mostly less than twice as long as broad; plants rare; fruit red, yellow, or orange 3 Petioles, at least some, with stalked red glands; teeth of leaves conspicuously tipped with reddish glands; plants known from Cache Co C. chrysocarpa Petioles lacking stalked red glands; teeth of leaves not conspicuously red tipped; plants known from Provo Canyon C. succulenta Crataegus chrysocarpa Ashe Yellow Haw- thorn. (C. rotundifolia Moench, not Lam.; C. doddsii Ramaley). Shrub or small tree with rounded crown, 2-4 m tall or more with thorns 1-5 cm long or more; leaf blades 1.8-7 cm long, 1-7.2 cm wide, orbicular to obo- vate, acute apically, acute to broadly obtuse basally, the margins sharply doubly serrate (the serrations red tipped) and commonly lobed as well; petioles often glandular (at least some); inflorescence more or less villous at anthesis; sepals lance-attenuate to triangu- lar, serrate; petals white, 6-8.5 mm long and about as broad; stamens 5-10; styles 2-4; fmit red to yellow-orange; "seeds" not pitted or deeply concave ventrally. Streamsides, re- ported from Cache Co. by Maguire (1937. Leafl. W. Bot. 2: 23-26); Alberta and Mani- toba south to Colorado and Nebraska. Crataegus douglasii Lindl. River Haw- thorn. Shrubs or small trees, with rounded crowns, mainly growing as thickets, 2.5-5 (7) m tall, with thorns 1.2-3.5 cm long; leaf blades 1.3-9.5 cm long, 0.8-5.8 cm wide, lan- ceolate to elliptic, oblanceolate, or obovate, acute to obtuse apically, cuneate basally, ser- rate to doubly serrate, seldom lobed; petioles often with a pair of raised sub-basal glands; inflorescence glabrous; sepals triangular-at- tenuate, entire or serrulate, 1.5-3.5 mm long; petals white, 3-7.8 mm long and about as broad or broader; stamens 10-20; styles nor- mally 5; fruit blackish, 6-12 mm thick. The species occurs from southeastern Alaska east to Michigan. Two rather well-defined varie- ties are present. 1. Petals 3-3.8 mm long, 2-4.2 mm wide; leaves slender, at least some 2-4 times longer than broad; fruit 6-8 mm thick when dried; plants of the Uinta Basin .... C. douglasii var. duchesnensis — Petals 4.5-8 mm long, 5-7.8 mm wide; leaves commonly less than twice longer than broad; fruit 8-12 mm thick when dried C. douglasii var. rivularis Var. duchesnensis Welsh var. nov. Similis Crataego douglasii var. rivularis sed differt in petalis breviore et angustiore foliis angustiore et pomis parviore. Type (in fruit). USA. Utah. Duchesne Co., Duchesne River valley, ca 24 Km northwest of Duchesne, along Utah Highway 35, Welsh, Atwood, and Moore 10928, 10 September 1970 (Holotype BRY). Additional specimens examined: Utah. Duch- esne Co., Duchesne River valley, Erdman 2516, 17 August 1965; do. Rock Creek, 12 mi WNW of Moimtain Home, Hansen s.n., 14 June 1976; do, 13 mi WNW of Mountain Home, Hansen s.n., 15 June 1976; do, T.IN., R.6W., Sec. 15, Hansen s.n., 14 June 1976; do, 5 mi NW of Hanna, Erdman 2522, 17 Au- gust 1965; Red Creek, ca 5 miles north of Fruitland, Brotherson 508, 21 June 1965. The Duchesne Hawthorn has long been recog- nized. It occurs along stream courses at 1800 to 2450 m in Duchesne and Uintah (?) coim- ties; endemic; 8(ii). Var. rivularis (Nutt.) Sarg. River Haw- thorn. (C. \ivularis Nutt. in T. & G.). Ter- races, flood plains, alluvial fans and canal banks and other moist sites at 1370 to 2135 m March 1982 Welsh: Utah Flora, Rosaceae 11 in Box Elder, Daggett, Juab, Millard, Piute, Salt Lake, San Juan, Sanpete, Sevier, Summit, and Utah counties; Wyoming and Idaho (?) south to Arizona and New Mexico; 34(iv). Crataegus monogyna Jacq. One-seeded Hawthorn. [C. oxijacantha var. monogyna (Jacq.) Loud.; C. oxyacantha L. nom. illeg.]. Shrub or small tree to 6 m tall, with thorns to 2 cm long; leaf blades 1.5-5.5 cm long and about as wide, broadly ovate to orbicular in outline, deeply 3- to 7-lobed, obtuse to trun- cate basally; petioles lacking glands; in- florescences glabrous to glabrate; sepals broadly triangular, 1.5-2.5 mm long; petals pink or white, 3-4 mm long, 4-5 mm wide; stamens commonly 20; styles 1 (2); fruit red- dish or orange, 5.5-8 mm thick. Cultivated ornamental in Cache, Davis, Salt Lake, Utah, Weber, and perhaps in other counties, escap- ing in some; introduced from the Old World; 1 (0). Crataegus succulenta Schrad. ex Link. Red Hawthorn. Shrubs or small trees, mainly 2-4 m tall with thorns to 4.5 cm long; leaf blades 1.8-7 cm long, 1-4.5 cm broad, elliptic to obovate, serrate to doubly serrate and often lobed; attenuate to cuneate basally, abruptly acuminate apically; petioles lacking glands inflorescence sparingly villous to glabrate petals white 5-7 mm long, 5.8-7.7 mm wide stamens 10-20; styles 2-4; fruit red, 7-12 mm thick when dry. Indigenous in riparian habitats, mainly in Provo Canyon, Utah Co.; Colorado eastward to Pennsylvania and southeastern Canada; 3(i). Note: Crataegus mollis (T. & G.) Scheele and/or C. crus-gallii L. and perhaps other species of hawthorn oc- cur in cultivation in Utah. The extent is not known at present. Cydonia Mill. Small trees, unarmed; leaves alternate, simple, entire; stipules foliose, glandular mar- gined, deciduous; flowers perfect, regular, solitary, terminal on leafy shoots; hypan- thium short; sepals 5, persistent; petals 5, white or pale pink; stamens 15-20; pistils 1, the ovary inferior, commonly 5-loculed; styles 5; frviit a tomentose pome. Cydonia oblonga Mill. Quince. {Pynis cy- donia L.; C. vulgaris Pers.). Trees to 6 m tall; leaves petiolate, the blades 1.1-9.5 cm long. 0.8-7 cm wide, ovate to ovate-oblong, vil- lous-tomentose beneath, tomentose above when young, becoming glabrate; flowers soli- tary; sepals foliose, 6-9 mm long, tomentose (especially within), glandular margined; pet- als 13-25 mm long, 8-17 mm wide, obovate to obcordate, white to pale pink; fruit 6-10 cm in diameter, yellow, densely tomentose, fragrant, broadly pyriform. Cultivated orna- mental and botanical curiosity in Utah and possibly other countries; introduced from the Middle East; 2 (0). Dryas L. Shrubs or subshrubs, with stoloniferous branches; leaves alternate, simple, crenate to entire, sometimes incised at the base, ever- green; stipules narrowly lanceolate, adnate to the petiole, persistent; flowers perfect (rarely imperfect), regular, solitary; hypanthium sau- cer shaped, with an internal glandular disk; sepals 8-10, persistent; petals 8-10; stamens numerous; pistils numerous, distinct, the ovaries superior, each 1-loculed; styles 1 per pistil, much elongated in fruit; fruit an ach- ene with a long-plumose style. HuLTEN, E. 1959. Studies in the genus Dryas. Svensk Bot. Tidskr. 5: 507-542. PoRSiLD, A. E. 1947. The genus Dryas in North America. Canad. Field-Naturalist 61: 175-192. Dryas octopetala L. Mat-forming shrubs; leaves petiolate, the petioles glabrous to spar- ingly villous and often glandular; leaf blades mostly 1-4 cm long, 0.3-1 cm broad, lanceo- late to lance-oblong, obtuse apically, obtuse to subcordate basally, crenate, green and glabrous to pubescent above, tomentose be- low, commonly with stipitate glands on the midrib below, often revolute; scapes 1-11 cm long, tomentose and stipitate-glandular; pet- als white (fading yellowish) or rarely yellow- ish, 9-15 mm long; staminal filaments glabrous; styles plumose, in fruit to 4 cm long. Moraines, slopes, and ridge crests in al- pine tundra and meadows at 3500-3965 m in Daggett, Duchesne, Summit, and Uintah counties; widespread in northern North America; circumboreal. Our material is as- signable to var. hookeriana (Juz.) Breitung (D. hookeriana Juz.; D. octopetala ssp. hook- eriana (Juz.) Hulten]; 9 (0). 12 Great Basin Naturalist Vol. 42, No. 1 Exochorda Lindl. Shrubs, deciduous, unarmed; leaves alter- nate, simple, entire or serrate; stipules none; flowers more or less imperfect (polygamo- dioecious), borne in terminal racemes; hypan- thium flaring with a broad disk internally; sepals 5; petals 5, white; stamens 15-25; pis- tils 5, connate except for the 5 free styles, the ovary superior, 5-loculed; fruit a bony capsule. Exochorda racemosa (Lindl.) Rehder Pearl Bush. {Amelanchier racemosa Lindl.; E. gran- diflora Hook.). Slender shrubs with spreading crowns, to 2.5 m tall or more, the herbage glabrous; leaf blades 1.2-6.5 cm long, 0.5-3.5 cm wide, elliptic to oblong or obovate, cu- neate basally, mucronate apically, entire or some serrate in the upper half; racemes 3- to 10-flowered; flowers very showy; sepals 1-2 mm long, broadly rounded, erose apically, chartaceous; petals 10-17 mm long. In- troduced ornamental in Utah County and in other low elevation urban areas; native to Asia; 2 (0). Fallugia Endl. Shrubs, deciduous, unarmed; leaves alter- nate, pinnately dissected; stipules adnate to the petiole, triangular-subulate, persistent; flowers mainly perfect, regular, terminal and solitary or in few-flowered cymes; hypan- thium hemispheric, persistent, hairy within; sepals 5, alternating with slender bractlets; petals 5, white; stamens numerous; pistils nu- merous, the ovaries superior, of 1 carpel each; style terminal; fruit an achene, tipped by the plumose style. Fallugia paradoxa (D. Don) Endl. Apache Plume. (Sieversia paradoxa D. Don). Shrubs to 1.5 m tall, the herbage stellate hairy, the bark scaly; leaves mainly 4-16 mm long, cuneate-flabellate, 3- to 5-lobed, green and lepidote above, rusty-lepidote beneath; pedi- cels 2-18 mm long; sepals 4-7 (11) mm long, broadly ovate, abruptly acuminate-cuspidate apically; petals 11-14 mm long, 8-15 mm wide, white; pistils numerous; styles plumose, 2-4 cm long in fruit. Wash bottoms in mixed desert shrub and pinyon-juniper communities at 940 to 2290 m in Garfield, Iron, Kane, San Juan, Washington, and Wayne counties; Ne- vada and California east to Texas and south to Mexico; 32 (vii). Fragaria L. Herbaceous, rosulate perennials, com- monly stoloniferous; leaves compound, with 3 serrate leaflets; stipules adnate to base of elongate petiole; flowers more or less imper- fect (polygamo-dioecious), solitary, or in sea- pose cymes; hypanthium widely spreading; sepals 5, alternating with bractlets; petals 5, white or pinkish; stamens 20, sometimes abortive; pistils numerous, on a pulpy recep- tacle, superior; fruit of achenes, on a fleshy accessory receptacle. Rydberg, p. a. 1908. Fragaria. N. Amer. Fl. 2: 356-365. 1. Petioles spreading-hairy; terminal tooth of leaflets relatively well developed, commonly surpassing the adjacent lateral teeth; inflorescence usually as long as or longer than the leaves p vesca Petioles with hairs ascending to appressed-ascending; terminal tooth of leaflets small, commonly surpassed by the adjacent lateral teeth; inflorescence usually shorter than the leaves p. virginiana Fragaria vesca L. Starvling Strawberry. Stoloniferous herbs, with stems, petioles, and peduncles pubescent with slender spreading to somewhat ascending hairs; petioles 0.8-17.5 cm long (rarely longer); leaflets 3, the terminal one 1.3-6.5 cm long, 1-4.2 cm wide, thin, elliptic to oblong or obovate, coarsely serrate, silky pilose, subsessile or in- distinctly petiolulate; scapes ultimately equaling or surpassing the leaves; cymes 3- to 15-flowered; sepals 3.5-7.2 mm long, acumi- nate to caudate; bracteoles 1.6-5.8 mm long, often bilobed; petals 5-10.5 mm long, white or pinkish; fruit to 1 cm thick, succulent, and palatable. Stream banks, terraces, and slopes, broad-leaved deciduous and coniferous woods and brushlands at 1800 to 3200 m in Box El- der, Cache, Carbon, Davis, Duchesne, Salt March 1982 Welsh: Utah Flora, Rosaceae 13 Lake, San Juan, Sanpete, Summit, Tooele, Utah, and Weber counties; British Columbia and Alberta south to California and New Mexico. Our material is referable to var. bracteata (Heller) R.J. Davis [F. bracteata Heller; F. vesca ssp. bracteata (Heller) Staudt.; F. helleri Holz.]; 34 (ii). Fragaria virginiana Duchesne. Mountain Strawberry. Stoloniferous herbs with stems, petioles, and pedimcles with appressed to as- cending hairs; petioles 2-15 cm long; leaflets 3, the terminal one 1.1-4.4 cm long, 0.5-2.2 cm wide, thickish, obovate to elliptic, coarse- ly serrate, silky pilose to glabrate, commonly petiolulate; scapes shorter than to surpassing the leaves; cymes 2- to 12-flowered; sepals 3.1-6.5 mm long; bracteoles 1.8-4.5 mm long, not or seldom bilobed; petals 3.5-10 mm long, white or rarely pinkish; fruit to 1 cm thick or more, succulent and palatable. Meadows, deciduous and coniferous woods at 2280 to 3300 m in Beaver, Duchesne, Emery, Garfield, Iron, Kane, Piute, Sanpete, Summit, Tooele, Utah, and Wayne counties; Alaska east to Northwest Territories and south to Colorado and California. Our material has been treated as belonging to two rather weak and intergrading varieties. The phase with large petals (i.e., more than 6 mm long) is supposedly more densely pubescent with spreading hairs. That phase is known as var. platypetala (Rydb.) Hall (F. platypetala Rydb.). It apparently intergrades completely with the small-flowered supposedly scantily pubescent phase with appressed hairs, known as var. glauca Wats. [F. vesca var. americana Rydb., not T.C. Porter; F. glauca (Wats.) Rydb.; F. virginiana ssp. glauca (Wats.) Staudt.]. In a broad sense, as herein inter- preted, all of our material is best referred to a single taxon. The oldest available epithet appears to be var. glauca Wats. Geum L. Perennial rhizomatous herbs; leaves alter- nate or opposite or mainly basal, pinnatifid to lyrate-pinnatifid; stipules foliose (at least on cauline leaves); flowers perfect, regular, solitary or in open cymes; hypanthium cam- panulate to saucer shaped; sepals 5, per- sistent, alternating with 5 bractlets; petals 5, usually yellow (sometimes pinkish or pur- plish); stamens numerous; pistils numerous, the ovaries superior, each 1-carpellate; style straight to bent or strongly geniculate and jointed, in some elongate in fruit and in some then with a deciduous terminal segment, in others plumose and persistent; fruit an achene. 1. Stems decidedly leafy; plants often more than 3.5 dm tall; sepals reflexed at anthesis; styles strongly geniculate and jointed, the persistent base hooked apically 2 — Stems subscapose; plants commonly less than 3.5 dm tall; sepals ascending to erect at anthesis; styles neither geniculate nor jointed 4 2(1). Persistent style base glandular-pubescent; terminal segment of basal leaves much larger than the lateral lobes, mostly rounded or subcordate at base; our common meadow and woodland Geurn G. macrophyllum — Persistent style base glabrous or hirsute, not glandular; terminal segment of basal leaves only somewhat larger than the lateral lobes, cuneate at base; plants uncommon 3 3(2). Petals equal to or shorter than the sepals; receptacle pubescent with coarse hairs; stem leaves with lobes or leaflets often about as broad as long, tapering to a rounded or acute apex; achenes about 70 G. urbanum — Petals longer than the sepals; receptacle minutely hairy; stem leaves with lobes or leaflets distinctly longer than broad and mostly tapering to an acute apex; achenes 200 or more G. aleppicum 14 4(1). Great Basin Naturalist Vol. 42, No. 1 Cauline leaves opposite; petals white, pink, or only yellow tinged, erect or convergent; styles much elongate and plumose in fruit G. triflorum Cauline leaves alternate; petals yellow, spreading; styles about as long as the achene, glabrous q rossii Geum allepicum Jacq. Erect Avens. [G. canadense Murr., not Jacq.; G. strictum Ait.; G. allepicum var. strictum (Ait.) Fern.; G. al- lepicum ssp. strictum (Ait.) Clausen]. Plants shortly rhizomatous, 4.5-8 (10) dm tall, the stems and petioles spreading-hirsute; basal leaves 8-23 cm long, lyrate-pinnatifid, main lobes 5-9, all cuneate-obovate, strongly cleft and toothed, the terminal lobe larger but sim- ilarly shaped; cauline leaves several; flowers 2 to several; sepals soon reflexed, 4-8 mm long; petals yellow, spreading, about equal- ing the sepals; stamens 60 or more; achenes 3-4.5 mm long, tipped by persistent style; style strongly geniculate above the middle, the lower segment hirsute to glabrous, not glandular near the base, persistent and hooked apically. Wet to dryish meadows at 1400 to 2300 m in Grand, Summit, and Utah counties; widespread in North America and Eurasia; 3 (0). Geum macrophyllum Willd. Large-leaved Avens. [G. urbanum ssp. oregonense Schintz; G. oregonense (Schintzl.) Rydb.; G. macro- phyllum var. njdhergii Farw.]. Plants shortly rhizomatous, 2.3-11.5 dm tall, the stems erect, with spreading hairs; basal leaves 4-28 cm long or more, long-petiolate, the leaflets 9-25 or more, the apical lobe 2.5-13 cm long, 2-15 cm wide, acute to truncate or sub- cordate basally, dentate and conspicuously lobed, glabrate to glabrous above, hairy along the veins beneath; cauline leaves numerous; sepals 2.5-4.8 mm long, reflexed at anthesis; bracteoles 0.5-3.5 mm long, linear to lanceo- late or lacking; petals yellow, 3-5.8 mm long; styles elongate with an s-shaped curve above the middle, glabrous or hairy above the bend, glandular (often sparingly so, or the glandular hairs deciduous) below the bend, hooked at apex, the apical section deciduous. Aspen, spruce-fir, birch-willow, and grass-sedge communities at 1280 to 2750 m in Box Elder, Cache, Carbon, Duchesne, Emery, Garfield, Kane, Piute, Salt Lake, Sanpete, Summit, Tooele, Uintah, Utah, Wasatch, and Wash- ington counties; widespread in North Ameri- ca; Asia. Our materials, as described above, belong to a poorly differentiated variety, the var. perincisum (Rydb.) Raup [G. perincisum Rydb.; G. macrophyllum ssp. perincisum (Rydb.) Hulten]; 40 (vii). Geum rossii (R. Br.) Ser. Ross Avens. [Sie- versia rossii R. Br.; Acomastylis rossii (R. Br.) Greene]. Herbs, shortly rhizomatous, the rhi- zomes and stem bases with a persistent thatch of marcescent petioles and stems; stems 0.3-1.8 dm tall, erect, often with 1-3 greatly reduced leaves below the in flores- cence; basal leaves 2.2-12 cm long, short-pe- tiolate, pinnatifid, with 15-31 entire to sever- al-toothed or -lobed lateral divisions, the apical lobes similar to the lateral ones except smaller, glabrous to pubescent along veins below, ciliate; sepals 2.9-5.1 mm long, ovate to ovate-lanceolate; bracteoles 2-3.9 mm long, lanceolate; petals yellow, 6.4-8.6 mm long; styles straight, elongate, glabrous ex- cept at base, erect. Alpine meadows, rock stripes, and talus slopes at 3050 to 3400 m in Daggett, Duchesne, Juab, Piute, Salt Lake, Summit, Uintah, Utah, and Wayne counties; Alaska and Yukon south to Oregon, Nevada, Arizona, and New Mexico; Asia. The speci- mens from Utah are referable to var. turhina- tum (Rydb.) C. L. Hitchc. [Potentilla nivalis Torr., not Lapeyr.; G. turbinatum Rydb.; Sie- versia turbinata (Rydb.) Greene; Acomastylis turbinata (Rydb.) Greene; G. sericeum Greene; S. sericea (Greene) Greene], which is only weakly separable from the larger typical phase that occurs in the arctic; 14 (ii). Geum triflorum Pursh Purple Avens; Old Man's Beard. [Sieversia triflora (Pursh) R. Br.; Erythrocoma triflora (Pursh) Greene; G. cilia- tum var. triflorum (Pursh) Jeps.]. Plants short- ly rhizomatous, clothed basally with per- sistent leaf bases; stems 1-4 din tall, with 1-2 pairs of opposite leaves; basal leaves 2-18.5 cm long, pinnatifid, with mostly 15-31 cleft to lobed divisions, puberulent to pilose; flow- ers 1-7 (9); sepals showy, reddish to pink or purplish, 7-10 mm long; bracteoles linear to narrowly elliptic; petals yellowish to pink or suffused with crimson, 7-11.2 mm long; styles straight to moderately geniculate, March 1982 Welsh: Utah Flora, Rosaceae 15 plumose, 2-4 cm long at maturity. Oak- sagebrush, aspen, aspen-fir, and mixed conifer communities, often in meadows, at 1980 to 3500 m in Beaver, Daggett, Garfield, Iron, Sanpete, Sevier, Summit, and Wayne coun- ties; British Columbia east to Newfoundland and south to Nevada, Arizona, New Mexico, Nebraska, and Illinois. Our specimens are as- signable to the weakly differentiated var. ciliatum (Pursh) Fassett [G. ciliatum Pursh; S. ciliata (Pursh) D. Don; Erythrocoma ciliata (Pursh) Greene]; 20 (v). Geum urbanum L. Plants shortly rhizo- matous, (3) 6-10 dm tall, the stems and pe- tioles spreading-hirsute; basal leaves mainly 2-10 cm long, pinnatifid, the main lobes 3-5 (10), cuneate-obovate, rounded to acute at the apex, the terminal one rhombic-ovate and slightly larger than the lateral ones; cauline leaves several to many; sepals spreading to reflexed, 4-7 mm long; petals yellow, spread- ing, somewhat shorter than the sepals; ach- enes 3-6 mm long, tipped by the persistent style; style strongly geniculate above the middle. Disturbed sites, reported from Salt Lake Coimty (Amow 5546, UT), adventive from Eurasia; 0 (0). HoLODiscus Maxim. Nom. Cons. Shrubs unarmed; leaves simple, alternate, toothed, deciduous; stipules lacking; flowers perfect, regular, each closely subtended by 1-3 bracteoles, numerous, borne in panicles; hypanthium saucer shaped, lined with a disk; sepals 5, persistent; petals 5, white to cream or less commonly pinkish; stamens about 20; pistils 5, the ovaries superior, each 1 -carpel- late; styles terminal, persistent; fruit a short- stipitate, villous achene. Ley, a. 1943. A taxonomic revision of the Genus Holodiscus (Rosaceae). Bull. Tor- rey Bot. Club 70: 275-288. Holodiscus dumosus (Nutt.) Heller Moun- tain Spray. [Spiraea dumosa Nutt. in T. & G.; S. discolor var. dumosa (Nutt.) Wats.; Schiz- onotus argenteus var. dumosus (Nutt.) Kuntze; H. discolor var. dumosa (Nutt.) Dip- pel; Schizonotus dumosus (Nutt.) Koehne; Schizonotus discolor var. dumosus (Nutt.) Rehder; Sericotheca dumosa (Nutt.) Rydb.; H. microphyllus Rydb., type from Alta; Serico- theca microphylla (Rydb.) Rydb.; H. discolor var. microphyllus (Rydb.) Jeps.; Sericotheca concolor Rydb.]. Shrubs, densely to intricately branched, mainly 0.5-1.5 m tall; main foliage leaves on spur branches, the blades 0.5-3.2 cm long, 0.2-2.3 cm wide, obovate to oblan- ceolate or elliptic, cuneate basally, promi- nently toothed or lobed, villous to glabrate or glabrous on one or both surfaces, pale be- neath; inflorescence 3-15 cm long; sepals 1.3-1.7 mm long, villous, sometimes pinkish; petals 1.9-2.2 mm long, white, cream, or pinkish; achenes somewhat flattened, villous- hirsute. Ubiquitous in numerous small plant communities, especially common on rock outcrops, slickrock plateau margins, and at bases of cliffs, or in talus slopes at 1280 to 3550 m in Beaver, Box Elder, Carbon, Davis, Duchesne, Emery, Garfield, Uintah, Utah, Washington, and Weber counties; Oregon east to Wyoming and south to California, Ne- vada, Arizona, and Mew Mexico. Attempts at recognition of infraspecific taxa are fraught with difficulties that are likely ecological rather than genetic reflections. A conserva- tive approach is indicated; 60 (xii). IVESIA T. & G. Perennial herbs, from a caudex; leaves pin- nately compound, alternate or primarily bas- al; stipules of cauline leaves foliose; flowers perfect, regular, borne in compact to open cymes; sepals 5, alternating with 5 brac- teoles; hypanthium saucer to cup shaped, lined with a disk; petals 5, yellow or white; stamens 5 or 20; pistils 1-15, the receptacle hairy, the ovaries superior; style subterminal; fruit of achenes. Keck, D. D. 1938. Revision of Horkelia and Ivesia. Lloydia 1: 75-142. 1. Leaflets usually fewer than 20; petals white to cream; plants of western Tooele Qq I. baileyi — Leaflets commonly 22-80 or more; petals white or yellow; plants of various distribution ^ 16 2(1). 3(2). 4(2). Great Basin Naturalist Vol. 42, No. 1 Petals white; stems more or less radiate-decumbent 3 Petals yellow; stems erect or ascending 4 Stamens 5; hypanthium cup shaped; plants of Utah, Salt Lake, and Summit counties /. utahensis Stamens 20; hypanthium saucer shaped; plants of Beaver, Garfield, Sevier, and Tooele counties J. kingii Hypanthium cup shaped; flowers in dense, congested cymes; plants widespread in central and northern Utah, our most common Ivesia /. gordonii Hypanthium saucer shaped; flowers in open cymes; plants of Beaver, Garfield, Kane, and Washington counties /. sabulosa Ivesia baileyi Wats. Plants 0.4-2.5 dm tall, from a woody caudex clothed with persistent leaf bases; herbage glandular-pubescent; bas- al leaves 3.5-12 cm long; leaflets 10-20, 3-15 mm long, parted to divided; hypanthium disklike; sepals 2.5-3.2 mm long, ovate- lanceolate; bracteoles lance-oblong to ovate; petals white or cream, about equaling the sepals; stamens 5; pistils 3-7; styles glandu- lar; achenes 1.6-2.3 mm long. Mountain slopes at 1700 to 3100 m in Tooele Co. (Deep Creek Mts.); eastern Nevada and adja- cent Utah. Out material belongs to var. se- tosa Wats. [Horkelia baileyi var. setosa (Wats.) Rydb.; /. setosa (Wats.) Rydb.; /. bail- eyi ssp. setosa (Wats.) Keck]; 1(0). Ivesia gordonii (Hook.) T. & G. Gordon Ivesia. [Horkelia gordonii Hook.; Potentilla gordonii (Hook.) Greene]. Plants erect, 7-30 cm tall, from a thick woody caudex clothed with persistent leaf bases; herbage pu- berulent, glandular-puberulent, or glabrous; basal leaves 1-25 cm long; leaflets 20-50 or more, 2-17 mm long, divided to base; cymes congested, many-flowered, 1 or few or sever- al; hypanthium campanulate; sepals 2-6.5 mm long, erect at anthesis, triangular-sub- ulate; bracteoles 1.2-2.8 mm long; stamens 5; pistils 1-6; style not glandular; achenes 1.7-2.1 mm long. Ponderosa pine, spruce-fir, and mixed conifer woods and upwards in al- pine sites, often in rocky meadows, at 2050 to 3660 m in Beaver, Davis, Duchesne, Morgan, Piute, Salt Lake, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, and Weber counties; Washington east to Montana and south to California and Colorado; 35(iv). Ivesia kingii Wats. King Ivesia. [Potentilla kingii (Wats.) Greene; Horkelia kingii (Wats.) Rydb.; P. eremica Gov.; H. eremica (Gov.) Rydb.; P. kingii var. incerta Jones; /. halo- phila Heller]. Plants decumbent, the stems 5-22 cm long, radiating from a thickened caudex clothed with blackish persistent leaf bases; herbage glabrous or pubescent, not glandular; basal leaves 0.5-12 cm long or more; leaflets 24-60 or more, 1-6 mm long, entire or ternately divided; cymes dichoto- mously divided, open; hypanthium saucer shaped; sepals 2.5-3.1 mm long, spreading at anthesis, lance-attenuate; bracteoles 1.3-1.9 mm long, ovate-lanceolate; petals white, 3-4.2 mm long, 2.5-3 mm wide, claws; sta- mens 20; pistils 2-9; styles not glandular; achenes 1.8-2.2 mm long. Saline meadows and pans at 1700 to 2380 m in Beaver, Gar- field, Sevier, and Tooele counties; Nevada and California. The plants tend to blend with the pale substrate of the saline pans, perhaps accounting for the paucity of our records from Utah; 8(v). Ivesia sabulosa (Jones) Keck Sevier Ivesia. [Potentilla sabulosa Jones, types from head of Sevier River, "25 miles south of Panguitch"; Comarella sabulosa (Jones) Rydb.; Horkelia sabulosa (Jones) L. O. Williams; H. mutabilis Brandegee; /. mutabilis (Brandegee) Rydb.]. Plants erect, the stems 10-42 (50) cm tall, from a woody caudex clothed with persistent leaf bases; herbage glabrous to villous and glandular; basal leaves 3.5-23 (30) cm long, the petioles often suffused red purple; leaflets 30-80, paired, 1-13 mm long, usually divided to the base; cymes branched, open; hypan- thium saucer shaped; sepals 3.3-5.5 mm long, triangular-acuminate, spreading at anthesis; bracteoles 1.3-2.7 mm long, lanceolate; pet- als yellow, 2.1-3 mm long, 0.3-0.5 mm wide; stamens 5; pistils 1-5; styles glabrous or nearly so; achenes 1.7-2.2 mm long. March 1982 Welsh: Utah Flora, Rosaceae 17 Sagebrush, pinyon-juniper, pygmy sagebrush, ponderosa pine and spruce communities, commonly on Hmestone at 2050 to 2750 m in Beaver, Garfield, Kane, and Washington counties; Arizona (?) and Nevada. The type locality of I. sabulosa is recondite. Jones states (see Leafl. West. Bot. 10: 216. 1965) that "... And 4 miles below Ranch I got 6031 to 33f." The type of I. sabulosa is his number 6032. Jones further states that "Ranch is the name of the post office at the head of the Sevier and serves a little farming area there." Presumably "Ranch" and the collecting site are both in Garfield County; 7(iv). Ivesia utahensis Wats. Utah Ivesia. [Poten- tilla utahensis (Wats.) Greene; Horkelia uta- hensis (Wats.) Rydb.] Plants decumbent or as- cending, the stems radiating from a thickened caudex clothed with brownish leaf bases; her- bage glandular-viscid or glandular-pubescent; basal leaves 1.2-9 cm long; leaflets 30-40, paired, 1-4 mm long, divided to base; cymes capitate, or in congested corymbs; hypan- thium campanulate; sepals 1.5-2.7 mm long, narrowly triangular; laracteoles 1-1.3 mm long, oblong; petals white, 1.8-2.7 mm long, 1-1.7 mm wide; stamens 5; pistils commonly 2; styles not glandular; achenes 1.7-1.9 mm long. Alpine timdra and krumholz commu- nities at 3200 to 3600 m in Salt Lake, Sum- mit, Utah, and Wasatch counties; endemic; 3(0). Kerria DC. Cultivated shrubs, unarmed, deciduous; leaves alternate, simple, doubly serrate; sti- pules lance-linear, deciduous; flowers perfect. regular, solitary, terminating short lateral branchlets of the current year; hypanthium short, lined by a disk; sepals 5, small, entire; petals 5, yellow; stamens numerous; pistils 5-8, the ovaries superior; style slender; fruit an achene (seldom produced). Kerria japonica (L.) DC. Japanese Kerria. {Rubus japonicus L.) Shrubs with slender green arching branches; leaves 1-7 cm long, 0.7-3.5 cm wide, ovate to lanceolate, long- acuminate, bright green; pedicels 0.5-2.5 cm long; sepals about 3.5 mm long, broad-ovate, membranous; petals yellow, 12-20 mm long; achenes 4-5 mm long. Widely cultivated or- namental, persisting but not spreading, ob- served in Box Elder, Cache, Davis, Salt Lake, Utah, and Weber counties; introduced from Japan and China. Double flowered phases are common; 2(i). Malus Mill. Trees with mainly unarmed branches; leaves alternate, simple, not or rarely lobed; stipules linear, caducous; flowers perfect, regular, borne in umbels; hypanthium short; sepals 5, persistent or deciduous; petals 5, white or pink; stamens usually 15-50; pistil 1, the ovary inferior, usually 5-loculed; styles 2-5, connate at the base; fruit variously shaped, applelike, the flesh usually lacking stone cells. Note: The apples are all in- troduced into our flora. Mostly they are culti- vated and persist following cultivation, but they escape and are established widely in Utah, especially the pomological varieties. The following key is tentative; our cultivars often represent hybrid derivatives involving two to several of the species types. Other taxa are probably present in the state. 1. 2(1). 3(2). Leaves on elongated shoots lobed or notched; a crabapple M. ioensis Leaves on elongated shoots neither lobed nor notched 2 Mature leaves glabrous on both surfaces, serrate to crenate-serrate, but not sharply so; fruit mainly 1.5-2.5 cm in diameter; a crabapple M. sylvestris Mature leaves more or less tomentose on one or both surfaces, or if glabrous, then the pedicels very long and the fruit smaller ^ Pedicels 3-4.5 cm long, very slender; fruit mainly 0.8-1.2 cm in diameter; a , , M. hupehensis crabapple ^ Pedicels mainly shorter than 3 cm long, slender to moderately thickened; fruit size various 18 4(3). 5(4). Great Basin Naturalist Vol. 42, No. 1 Leaf margins sharply serrate; flowers ordinarily pink; flowering ornamental crabapple M. floribunda Leaf margins merely crenate-serrate to serrate; flowers various 5 Flowers commonly pink; fruit 0.8-L2 cm in diameter; a crabapple M. baccata Flowers commonly white within (pink sometimes on dorsal surface) M. pumila Malus baccata (L.) Borkh. Siberian Crab. [Pyrus baccata L.]. Small trees to 5 m, the branchlets glabrous; leaves ovate to oblong, 2-8 cm long, serrate, glabrous to puberulent on one or both sides; petioles 1.5-5 cm long; petals white or pink, 12-18 cm long; fruit 0.8-1.2 cm in diameter, the calyx deciduous. Cultivated ornamental tree in lower eleva- tion portions of Utah; introduced from Asia; 4(0). Malus floribunda Sieb. ex Van Houtte Showy Crab. [Pyrus floribunda Sieb.]. Small trees to 8 m, the branchlets pubescent; leaves ovate to oblong, 2-7 cm long, sharply serrate, usually tomentose on one or both sides; pe- tioles mostly 1.5-5 cm long; petals rose pink, 15-20 mm long or more; fruit mostly 0.5-1 cm in diameter. Cultivated ornamental trees in lower elevation portions of Utah; in- troduced from Japan; 5 (0). Malus hupehensis (Pamp.) Rehder Tea Crab. [Pyrus hupehensis Pamp.; M. theifera Rehder]. Small trees to 5 m tall, the branch- lets glabrous or essentially so; petioles 0.8-4 cm long; leaf blades oblong-elliptic to ovate, 2.5-7.8 cm long, minutely tomentulose on both sides at maturity; peduncles 3-4.5 cm long; sepals 4-6 mm long; petals 15-20 mm long, commonly white; fruit 0.8-1.2 cm in di- ameter. Cultivated ornamental trees of lower elevations in Utah; introduced from Asia; 7 (0). Malus ioensis (Wood) Britt. Iowa Crab. [Pyrus ioensis (Wood) Bailey]. Small trees to 9 m tall, the branchlets tomentose; leaves ovate to oblong, 2.5-10 cm long, tomentose on both sides at least when young; petals usu- ally white (sometimes pinkish), 12-25 mm long; fruit 2-3 cm in diameter. The Iowa crab is grown occasionally in lower elevation portions of Utah; introduced from the north central states; 4 (0). Malus pumila Mill. Common Apple. [M. domestica Borkh.]. Small to moderate trees to 10 m tall, the branchlets tomentose when young, becoming glabrous; leaves ovate to oblong or elliptic, 1.5-10 cm long, tomentose on one or both sides (even in age); petals usu- ally white within, often pink dorsally, 12-25 mm long; fruit mainly 2.5-12 cm in diame- ter, red, reddish purple, or yellow. This is the apple of commerce, and it is widely culti- vated in Utah; it persists and occurs as estab- lished trees throughout the state; introduced from Eurasia; 18 (ii). Malus sylvestris Mill. Crabapple. Small to moderate trees to 10 m tall, the branchlets glabrous or puberulent when young, often somewhat thorny; leaf blades 2-6 cm long, ovate to elliptic, crenate-serrate, acuminate or cuspidate; petals 8-20 mm long, white or pink; fruit mainly 1.5-2.5 cm in diameter, sour. Introduced cultivated trees, persisting and escaping in Utah; native to Eurasia; 6 (1). Peraphyllum Nutt. in T. & G. Shrubs, unarmed, deciduous; leaves alter- nate, simple, entire or nearly so; stipules ad- nate to petioles, triangular, minute, de- ciduous; flowers perfect, regular, solitary or few on lateral branchlets of the season; hy- panthium campanulate, disk lined; sepals 5, spreading to reflexed, persistent; petals 5, white or pink; stamens 15-20; pistil 1, the ovary inferior, 2- to 3-carpellate but falsely 4- to 6-loculed by intrusion of parietal septa; styles 2 or 3, the stigma capitate; fruit a fleshy apple-like pome. Peraphyllum ramosissimum Nutt. in Torr. & Gray. Squaw-apple. Shrubs 4-15 (20) dm tall, intricately branched; leaves alternate, mainly on short lateral spurs, 1.1-3.9 cm long, 0.4-0.9 cm wide, oblanceolate, abruptly acute, appressed puberulent (especially be- neath), entire or minutely serrulate; pedicels 4-13 mm long, with 1-3 caducous bractlets; sepals 2.9-4 mm long, triangular-acuminate, serrulate to entire; petals white to pink, 6.5-9 mm long, 5-8.5 mm wide; pomes 8-18 mm thick, yellow orange, the flavor bad when ripe. Oak-sagebrush, pinyon-juniper. March 1982 Welsh: Utah Flora, Rosaceae 19 mountain brush, and ponderosa pine commu- nities at 1500 to 2500 m in Beaver, Garfield, Grand, Juab, Kane, Millard, San Juan, San- pete, Sevier, and Washington counties; Ore- gon and Idaho south to California and Colo- rado. The fruit is attractive when ripe but the flavor is not agreeable. Perhaps, when cooked with sugar it might be better?; 20 (vi). Petrophytum (Nutt.) Rydb. Shrubs, prostrate and mat forming, con- forming to the rock substrate; leaves alter- nate, commonly appearing rosulate, simple, entire, stipules lacking; flowers perfect, regu- lar, borne in compact spikelike panicles on scapose, bracteate peduncles; pedicels with 1 or more bractlets; hypanthium cup shaped, lined with a disk; sepals 5, erect at anthesis; petals 5, white; stamens numerous; pistils commonly 5, the ovaries superior, each l-lo- culed; styles slender, exserted from the flow- er; fruit of usually 5 follicles. Petrophytum caespitosum (Nutt.) Rydb. Rock Spiraea. [Spiraea caespitosa Nutt. in T. & G.; Eriogyna caespitosa (Nutt.) Wats.; Luetkea caespitosa (Nutt.) Kuntze]. Mat- forming shrubs to 10 dm broad or more; leaves 3-17 mm long, 1.5-4.5 mm wide, spatulate to oblanceolate or obovate, pilose on one or botli surfaces, rarely almost or quite glabrous; peduncles 0.5-12 cm long, with bractlike leaves much reduced upwards; panicles spikelike, 0.5-2.5 cm long, often branched at base or with axillary panicles along axis of peduncle; pedicels 0.5-3 mm long, bracteolate; sepals 1.2-2.1 mm long. narrowly triangular; petals 1.3-2.5 mm long, 0.4-0.8 mm wide, white; fruit 1.5-2.1 mm long. On limestone or granitic outcrops or gravels from sagebrush upwards to spruce-fir communities and on sandstone (Entrada, Navajo, Kayenta, Cedar Mesa, etc.) often in hanging gardens at lower elevations (1000 to 2750 m) in Beaver, Cache, Grand, Juab, Kane, Millard, Utah, and Washington coun- ties; Oregon east to South Dakota and south to California, Arizona, New Mexico and Texas. This beautiful dwarf shrub flowers in late summer and autumn. Major variations involve the tendency to glabrous leaves of some Great Basin specimens, and a tendency to short peduncles in some materials from the hanging gardens of southeastern Utah. More materials are required to adequately assess the variation; 25 (vii). Physocarpus Maxim. Nom. Cons. Shrubs, unarmed, deciduous, with exfoliat- ing bark; leaves alternate, simple, palmately lobed and veined, usually with at least some stellate hairs; stipules membranous, de- ciduous; flowers perfect, regular, borne in terminal corymbs; hypanthium cup shaped, lined with a disk; sepals 5; petals 5, white or pink; stamens 20-40, inserted with petals at edge of disk; pistils 1-5, the ovaries superior and partially connate; styles slender, the stigmas capitate; fruit of one or more fol- licles, each several seeded. Howell, J. T. 1931. A Great Basin species of Physocarpus. Proc. Calif. Acad. Sci. IV. 20:129-134. 2(1). Pistil and style solitary; leaves less than 2 cm long; staminal filaments of two alternating and markedly unequal lengths P- alternans Pistils and styles 2 or 3, or the carpels connate below; leaves various but commonly over 2 cm long; staminal filaments subequal or somewhat unequal 2 Leaves mainly 0.7-2.5 cm long; mature carpels swollen, not flattened; plants evidently rare in Utah P- monogynus Leaves mainly 2-8 cm long; mature carpels flattened; plants common in south central to northern Utah P- malvaceus Physocarpus alternans (Jones) J. T. How- Heller] Shrubs commonly 4-12 dm tall, and ell Dwarf Ninebark. [Niellia monogyna var. about as broad; twigs ste late pubescent and alternaiis Jones; Opulaster alternans (Jones) sometimes glandular; bark shreddy on older 20 Great Basin Naturalist Vol. 42, No. 1 twigs; leaf blades 0.3-2 cm long, 0.3-2.2 cm wide, oval-ovate to ovate, cordate to sub- cordate basally, more or less 3-lobed, doubly crenate, pubescent to sparingly pubescent on both sides; inflorescence subumbellate, (1) 2- to 12 (17) -flowered; pedicels 2-10 mm long; hypanthium stellate-hairy; sepals 1.3-3.2 mm long, oval to suboblong; petals 1.8-3.2 mm long, 1.5-3 mm wide, white or suffused with red pink; follicle solitary, densely stellate, 4-5 mm long. Rock outcrops, ledges, and cliff faces in desert shrub, pinyon-juniper, oak, and ponderosa pine communities at 1980 to 2750 m in Daggett, Duchesne, Emery, Gar- field, Millard, San Juan, Utah, Wayne, and Washington counties; Idaho, Nevada, and Colorado. Jones (Zoe 4:38-44. 1893) dis- cussed the problems within the genus Phys- ocarpus then called Niellia. The basic prob- lems are still as Jones outlined them eight decades ago. The genus is still in need of a definitive revision; 14(iv). Physocarpus malvaceus (Greene) Kuntze Mallow-leaved Ninebark. [Niellia malvacea Greene; Opulaster malvaceus (Greene) Kuntze; N. monogyna var. malvacea (Greene) Jones; Spiraea opulifolia var. pauciflora T. & G.; Opulaster pubescens Rydb.; O. cordatus Rydb.j Shrubs, mainly 8-20 dm tall, rarely more, and often as broad; twigs glabrous to minutely stellate; bark shreddy on older branchlets; leaf blades (0.8) 2.2-8 cm long, (1.2) 2-8.2 cm wide, ovate to broadly ovate, cordate basally, 3-lobed, doubly crenate, glabrous above, stellate-pubescent to glabrous beneath; inflorescence corymbose, 5- to 30( -I- )-flowered; pedicels 0.7-2.3 cm long; hypanthium stellate-hairy; sepals 2.2-4.6 mm long, ovate to lance-oblong; pet- als 3.3-6.7 mm long, 1.5-4.8 mm wide, white; follicles paired, connate to the middle or above, substipitate, densely stellate, 4.9-6 mm long. Moist slopes and streamsides in mountain brush, aspen, and mixed conifer woodlands at 1600 to 3300 m in Cache, Emery, Garfleld, Juab, Millard, Salt Lake, Sanpete, Summit, Tooele, Utah, Wasatch, and Weber counties; British Columbia east to Alberta and south to Oregon, and Wyoming; 39(vii). Two other large-leaved ninebark spe- cies are known from cultivation in Utah. They are P. opulifolius (L.) Raf. {Spiraea opulifolia L.) and P. capitata (Pursh) Kuntze {Spiraea capitata Pursh). They both possess 3-5 glabrous pistils connate only at the base. They differ in that the leaves of P. opulifolius are commonly glabrous beneath and those of P. capitatus are ordinarily densely stellate be- neath. The extent of these species in cultiva- tion is not known. Physocarpus monogynus (Torr.) Coult. Mountain Ninebark. [Spiraea monogyna Torr.; Opulaster monogynus (Torr.) Kuntze]. Shrubs, mainly 4-20 dm tall; twigs glabrous to densely stellate; bark shreddy on older branchlets; leaf blades 0.5-2.8 cm long, 0.6-3.2 cm wide, ovate to orbicular, cordate basally, commonly 3-lobed, doubly crenate, glabrate or glabrous on both sides or some- times stellate hairy especially below; in- florescence corymbose, 9- to 25-flowered (sometimes more); pedicels 0.3-0.8 cm long; hypanthium stellate-hairy; sepals 2.3-3 mm long, ovate; petals 2.2-3.9 mm long, 2.6-3.5 mm wide, white; follicles paired, connate to the middle or above, substipitate, densely stellate; 3-4.5 mm long. Canyon bottoms and moist slopes in mountain brush, aspen and Douglas fir communities at 1650 to 2150 m in Carbon and Utah counties; South Dakota and Wyoming south to Texas and Arizona (?); 2 (i). Our material seems to flt well within the range of variation of materials from Colo- rado, New Mexico, and South Dakota. The plants are smaller in all features from the similar P. malvaceus (q.v.). POTENTILLA L. S. L. Welsh & B. C. Johnston Annual, biennial, or perennial herbs (a shrub in P. fruticosa); leaves alternate or bas- al, palmately or pinnately compound; sti- pules lanceolate to ovate, sometimes sheath- ing; flowers perfect, regular, borne solitary or in cymes; hypanthium saucer shaped to cup shaped; sepals 5, alternating with bractlets; petals 5, yellow to ochroleucous (fading white in some), broadly obovate and often emarginate; stamens 10-25; pistils numerous, on a hemispheric to conical receptacle, supe- rior; fruit of achenes, the styles terminally, medially, or basally attached, jointed, finally deciduous, glabrous or papillose. Note: This is a fairly difficult genus, with many inter- March 1982 Welsh: Utah Flora, Rosaceae 21 pretations, due partially to hybridization, I. Effect of varied environments on which obscures differences between taxa, and western north American plants. Car- because of different weight given to portions negie Inst. Washington Publ. 520: of the genus as providing basis of segregation 31-49. Potentilla gracilis and its allies, into several genera. We have resisted segre- op. cit., pp. 128-137. gation of the Utah species into the genera Ar- Lehmann, J. C. G. 1856. Revisio Potentilla- lentina, Drymocallis, and PentaphyUoides. rum Iconibus Illustrata. Verh. Kaiserl. These generic names will be used by some Leopold Akad. Vol. 23. suppl. authors nevertheless, and we have included Rydberg, P. A. 1898. Monograph of the the designations for them in the following North American Potentillae. Mem. key. These names, as svnonyms, are provided Dept. Bot. Columbia Univ^ No 2. in the body of the treatment. 1908. Potentilleae. N. Amer. Fl. Hitchcock, C. L. 1961. Rosaceae. In C. L. 22(%): 268-377. Hitchcock, A. Cronquist, M. Ownbey, 1922. Flora of the Rocky Mountains and J. W. Thompson. Vascular Plants of and Adjacent Plains. New York, by the the Pacific Northwest. Univ. Washing- author, ed. 2. ton Publ. Biol. 17(3): 89-194. Tidestrom, I. 1925. Flora of Utah and Ne- Keck, D. D. 1940. Potentilla glandulosa and vada. Contr. U.S. Natl. Herb. 25: its allies: taxonomy. In J. Clausen, D. 1-665. , r- . D Keck and W M. Hiesey. Environ- Wolf, T. 1908. Monographic der Gattung mental studies on the nature of species. Potentilla. Bibliotheca Botanica 16(71). 1 Plants woody shrubs; styles laterally attached to the ovary; ovaries and achenes hairy (PentaphyUoides Duham.) ^- Ftiticosa Plants herbaceous; styles basal, terminal, or lateral; ovaries and achenes glabrous 2(1) Plants strongly stoloniferous; styles laterally attached to the ovary; leaves evi- dently pinnate, strongly bicolored, silvery white beneath {Argentina Lam.) -^ ^ P. anserina Plants without stolons, or rarely somewhat stoloniferous; styles basal or terminal; leaves usually not evidently pinnate and strongly bicolored 3 3(2) Styles basally attached to the ovary, relatively short (ca 1 mm) and deltoid gradually tapering upward from the flaring base; plants medium to tall, with pinnate leaves and often flabellate leaflets {Drymocallis Lam.) 4 Styles terminal, sometimes short, but not deltoid, if thickened at base then abruptly tapered; plants prostrate, short, or tall; leaves pinnate to palmate ^ {Potentilla L., sens, str.) 4(3). Leaflets 9-11, usually dark green, deeply and sharply toothed, often with inter medium to pale green, larger; flowers yellow to white spersed smaller leaflets between paired ones; stems often stoloniferous, less than 30 cm tall; flowers creamy white; plants rare ^- /»««« Leaflets 5-9, usually lacking interspersed smaller leaflets; stems and leaves 5(4) Inflorescence short and dense, capitate, with erect branches; stem stout, often densely brown-villous; petals creamy white, about equaling the sepals; plants mostly 40 cm tall or more; plants uncommon P- arguta Inflorescence open, with diverging branches, or if dense then the stems neither stout nor villous; petals cream to yellow; plants mostly less than 40 cm . ,, 1 , P. glandulosa tall; plants common » 22 Great Basin Naturalist Vol. 42, No. 1 6(4). Stems branched at or below the middle, the basal leaves lacking and midstem leaves developed at anthesis; styles to 1 mm long, conspicuously thickened at base; plants annual or short-lived perennials, of damp disturbed sites 7 - Stems usually branched above the middle, the basal leaves persistent at an- thesis, the midstem leaves reduced; style 0.8-4.5 mm long, thickened at base or not; plants perennial 21 7(6). Leaves conspicuously white, tomentose beneath, small, palmate, dissected into narrow lobes; plants adventive, to be expected in Utah P. argentea L. - Leaves not tomentose, strigose and often glandular, palmate or pinnate 8 8(7). Leaves all apparently pinnately compound, with 5-7 leaflets; achenes with a thickened appendage on inner margin p paradoxa - Leaves (at least the uppermost) ternate or palmate, or if the lowermost ones pinnate then leaflets crowded and subpalmate; achenes without an appendage 9 9(8). Lowermost leaves crowded-pinnate or subpalmate, with 5-7 leaflets; stems and calyx without glandular or pustulate hairs ' p rivalis - Lower leaves ternate or palmate; stems and calyx glandular or hirsute with pustulate hairs in 10(9). Stems covered with glandular hairs; leaves all ternate; petals often shorter than the sepals P. biennis - Stems with pustulate, spreading, stiff-hirsute hairs, and sometimes also with few glandular hairs; leaves ternate to palmate; petals often equaling sepals P. norvegica 11(6). Style to 1 mm long, often thickened at base, relatively thick just below the stigma; leaves tomentose, at least below 22 - Style 1.2 mm long or more, usually over 1.5 mm, often narrow just below the stigma jg 12(11). Leaves ternate, or less commonly 5-foliolate, or subpalmate, snowy white- tomentose beneath, the margins usually plane; inflorescence open; calyx usually lacking glands; plants alpine 13 - Leaves pinnate, or less commonly subpalmate, olive greenish or yellowish- tomentose beneath, the margins usually revolute; inflorescence glomerate, ODmpact; calyx glandular; plants variously distributed P. pensylvanica 13(12). Leaves subpalmate, with 5 leaflets; petioles strigose and tomentose; styles conelike, conspicuously thickened and papillose at base 14 - Leaves ternate; petioles strigose and tomentose, or only tomentose; styles uni- formly thickened, or sometimes only somewhat thickened below the stigma, rarely thickened at base ' 15 14(13). Leaves somewhat subpalmate, the rachis very short (ca 5-15 percent of petiole length), conspicuously strigose on petioles and on both surfaces P. rubricaiilis - Leaves subpalmate to subpinnate, the rachis longer (ca 20-40 percent of petiole length), tomentose beneath P. nivea x P. pensylvanica var. paucijiiga 15(13). Petioles strigose and tomentose; pubes cence snow white to off-white; plants of- ten more than 10 cm tall, rare in the Deep Creek, La Sal, and Uinta "lO'^ntains Phookeriana - Petioles tomentose only; pubescence snow white; plants commonly less than 10 cm tall, rare, alpine, in La Sal Mountains p. nivea March 1982 Welsh: Utah Flora, Rosaceae 23 1 7 16(11). Leaves palmate to subpalmate, not pinnate ^ — Leaves pinnate 17(16). Upper leaf surfaces and calyx glandular, often also strigose; stems to 15 cm long (rarely more), prostrate-ascending ^^ Upper leaf surfaces and calyx glabrous, sericeous, or tomentose; stem length various, but usually over 15 cm long ^^ 18(17) Petioles to 5 cm long; leaflets usually bicolored, tomentose beneath, strigose (at least) above; sepals and bractlets relatively broad, usually deltoid; styles 1.6-2.4 mm long, sometimes clavate; plants widespread P. concinna Petioles usually over 5 cm long; leaflets strigose on both surfaces, not bi- colored; sepals and bractlets narrow, acuminate; styles 2.1-3 mm long, filiform ; p. multisecta 19(18) Leaves bicolored, tomentose beneath; leaflets 7-9, palmate or subpalmate, short-serrate below middle, ovate or obovate; style filiform, if thickened, then 1- Ui-i t-u^^^ P. piilcherrima slightly so at base r Leaves not bicolored, or only slightly so, if tomentose beneath then not densely so and the leaflets oblanceolate or dissected almost to midrib; leaflets 5-7, pal- mate to subpalmate, not toothed below the middle in some; style filiform or uniformly thickened 90(19) Leaflets toothed above the middle, cuneate, never bicolored, green to grayish green- anthers mostly 0.5-0.7 mm long; stems decumbent to ascendmg; p ants °, . u 1 • P- diversifolia alpine or subalpine ^ ■' - Leaflets toothed to below the middle, often bicolored and pale beneath; anthers 0.8-1.3 mm long; stems ascending to erect 21 21(20). Leaflets divided two-thirds or more to midrib into linear segments, or some segments more than 5 mm long, whitish or grayish beneath P. pectimsecta Leaflets divided to one-half the distance to midrib, the teeth les than 5 mm long, usually green or greenish on both surfaces P- gracilis 22(16) Leaflets tomentose, at least below, rarely strigose or glandular; stems usually ascending, 20-60 cm tall; leaflets sometimes confluent with rachis; plants of montane to subalpine ~' Leaflets seldom tomentose, or if so then sericeous, strigose or glandular; stems decumbent to ascending, 5-35 cm tall, or if taller then leaflets strigose and glandular; leaflets not confluent with rachis; plants of submontane, montane, ^^ or alpine sites *" 23(22) Calyx densely tomentose at anthesis, with dark bractlets much smaller than the lobes; leaflets tomentose above, not strongly bicolored, 9- to 19-toothed; stems leafy; plants of the Uinta Mts I . effusa Calyx sericeous at anthesis, the bractlets similarly colored and subequal to the lobes; leaflets various' above, often strongly bicolored, 13- to 37-toothed; ^^^^^^ .^^^ not leafy; plants widespread nppmna 24(22). Leaflets strigose and usually also glandular beneath sometimes also tomentose above- pedicels straight; inflorescence branches divaricate; plants of south ^ ', Tu u P. crinita central Utah Leaflets glabrous to sericeous, if strigose (as rarely) then plants with recurved pedicels; inflorescence open, but branches not divaricate; plants mainly ot ^^ northern Utah 2'* Great Basin Naturalist Vol. 42, No. 1 25(24). Pedicels recurved in fruit; lower leaflets conspicuously pinnately toothed; stems decumbent or spreading; plants rare, in wet meadows P. platensis - Pedicels straight or ascending in fruit; lower leaflets pinnately toothed or api- cally few toothed; stems decumbent to ascending; plants montane to alpine P. ovina Potentilla anserina L. Common Silver- weed. [Argentina anserina (L.) Rydb.; A. ar- gentea Rydb.; P. anserina var. grandis T. & G.; A. anserina var. grandis (T. & G.) Rydb.; P. egedii var. grandis (T. & G.) Howell; P. pa- cifica Howell; A. pacifica (Howell) Rydb.]. Perennial herbs, with long strawberrylike sto- lons; leaves 2-100 cm long or more, pin- nately compound with 5-25 main leaflets in- terspersed by smaller ones, the terminal leaflet 0.5-5.5 cm long, 0.3-2.6 cm wide, oval to oblong or oblanceolate to obovate, coarsely serrate, green and glabrous to pilose above, pale and villous over a tomentum be- neath; scapes 1.5-15 cm long or more, villous to densely so, leafless; sepals ovate, 3-10 mm long, pubescent to glabrous, erect, enlarging in fruit; petals yellow, 7.5-16 mm long; ach- enes 1.5-2 mm long. Meadows, lake shores, terraces, and floodplains, especially where wet part of the season, at 1300 to 2600 m in Carbon, Garfleld, Kane, Millard, Piute, Salt Lake, Sanpete, Sevier, and Utah counties; widespread in North America; circumboreal. The worldwide review by Rousi (Ann. Hot. Fenn. 2: 47-112. 1965) indicates that formal varieties are not warranted; 24 (ii). Potentilla arguta Pursh Acute Cinquefoil. (P. pensylvanica var. arguta (Pursh) Ser. in DC; P. agrimonioides var. arguta (Pursh) Farw.; Drymocallis arguta (Pursh) Rydb.; Geum agrimonioides Pursh; P. agrimonioides (Pursh) Farw., not Beib.; D. agrimonioides (Pursh) Rydb.; P. convallaria Rydb.; D. con- vallaria (Rydb.) Rydb.; P. arguta var. con- vallaria (Rydb.) T. Wolf; D. corymbosa Rydb.) Perennial, glandular-pubescent herbs, 2.5-6 (8) dm tall, from a caudex; basal leaves 6-30 mm long, pinnately compound with 5-11 leaflets, the terminal one 15-60 mm long (or more), 12-40 mm wide, oval to ellip- tic or obovate, doubly dentate to somewhat lobed, green and glandular-pubescent on both surfaces; flowers several to many, showy; sepals 4-8 mm long, lance-ovate, longer in fruit; bracteoles 2-6 mm long, ob- long to narrowly lanceolate; petals yellow to cream or white, mostly 5-8 mm long; recep- tacle sparsely hairy; achenes 1-1.5 mm long; styles basal, ca. 1.0 mm long, deltoid. Moun- tain brush, aspen, and spruce-fir commu- nities, often in meadows, at 1950 to 3360 m in Davis, Juab, Salt Lake, San Juan, Sanpete, Summit, Utah, Washington (?), and Weber counties; the species is widespread in North America. Our material is referable to var. convallaria (Rydb.) T. Wolf, and is transi- tional to P. glandulosa (q.v.); 14 (i). Potentilla biennis Greene Green Cinque- foil. [Tridophyllum bienne (Greene) Greene; P. lateriflora Rydb., type from Utah; P. kel- seyi Rydb.] Annual or biennial herbs, mostly 1-6 (7) dm tall, from taproots; leaves mainly cauline, palmately 3 (4) -foliolate, the termi- nal leaflet 1-5 cm long, 1-3 cm wide, obo- vate to oblanceolate, crenate-serrate, pu- bescent with spreading to appressed hairs and multicellular glandular ones; flowers sev- eral to numerous, inconspicuous; sepals most- ly 2-4 mm long, ovate to lance-ovate; brac- teoles 2-3 mm long, ovate-lanceolate to oblong; petals yellow, 1.5-3 mm long; ach- enes numerous, about 1 mm long; styles ter- minal, ca. 1.0 mm long, basally thickened. Meadows, streamsides, springs, and seeps at 1525 to 2324 m in Beaver, Duchesne, Gar- field, Iron, Salt Lake, Sevier, Wasatch, and Washington counties; widespread in western North America; 7 (v). Potentilla concinna Richards. Pretty Cin- quefoil. Perennial herbs, the stems decum- bent-spreading to ascending, 0.1-1 dm tall; leaves mainly basal, palmately to pinnately 5- to 7 (9) -foliolate, the terminal leaflet 0.3-3.8 cm long, 0.1-1 cm wide, obovate to oblanceolate, toothed only at apex or along the length, often folded, not markedly bi- colored, pilose and tomentose beneath, pilose to glabrous (less commonly tomentose) above; cauline leaves 1 or 2; cymes (1) 2- to 7-flowered, the flowers showy; sepals 2.2-4.8 mm long, triangular-ovate; bracteoles 1.3-3.5 mm long, oblong to lanceolate; petals yellow, 2.7-7.3 mm long; achenes numerous, 1.6-2 March 1982 Welsh: Utah Flora, Rosaceae 25 mm long; style subapically attached, smooth times clavate. Three rather weak varieties or glandular, not basally thickened, some- are present in Utah, separable as follows: 2(1). Leaflets pinnately disposed, often 7, the lowermost often scattered or reduced; stems usually trailing, longer than basal leaves; pubescence translucent- brownish; plants of western Utah P. concinna var. proxima Leaflets palmately disposed, commonly 5-9; stems shorter than basal leaves; pubescence white ^ Leaflets toothed from the middle or below (at least some), commonly flat and green above; plants widespread -P. concinna var. modesta Leaflets toothed at apex only, rarely minutely crenate also, commonly folded, flie obscured upper surface greenish or not P. concinna var. bicrenata Var. bicrenata (Rydb.) Welsh & Johnston, comb. nov. [based on: Potentilla bicrenata Rydb. Bull. Torrey Bot. Club 23: 431. 1896]. Pinyon-juniper, grassland, sagebrush, pon- derosa pine, and spruce-fir communities at 2050 to 2875 m in Beaver, Duchesne, Emery, Garfield, San Juan, Sevier, and Wayne coun- ties; Colorado, Wyoming, and New Mexico; 14 (ii). Var. modesta (Rydb.) Welsh & Johnston, comb. nov. [based on: Potentilla modesta Rydb. N. Amer. Fl. 22: 331. 1908]. Sage- brush, meadow, aspen, spruce-fir, and Douglas flr communities at 2280 to 3480 m in Carbon, Emery, Garfield, Piute (type from Mt. Barrett near Marysvale), Sanpete, Sevier, Wasatch, and Wayne counties, endemic. The modesta phase of the pretty cinquefoil occurs generally above the range of var. bicrenata, with which it forms occasional intermediates. Specimens from Emery and Sanpete counties approach var. divisa Rydb. and P. gracilis; 18 Var. proxima (Rydb.) Welsh & Johnston comb. nov. [based on Potentilla proxima Rydb. N. Amer. Fl. 22(4): 339. 1908. (P. beanii Clokey; "P. quinquefolia" sens. Rydb.)]. Pinyon-juniper, sagebrush, pon- derosa pine, and spruce-fir communities at 2200 to 3000 m in Carbon, Garfield, Iron, Piute, and Wayne counties; Nevada; 34 (iii). Potentilla crinita Gray Hair-tuft Cinque- foil. Perennial herbs, the stems ascending to erect, 1.3-4.3 dm tall; leaves mainly basal, pinnately 7- to 13-foliolate, the terminal leaf- let 0.6-3 cm long, 0.2-0.9 cm wide, elliptic- oblong to narrowly oblanceolate, toothed at the apex only (somewhat below on some leaves), not markedly bicolored, pilose and sometimes slightly tomentose beneath, stri- gose-pilose to glabrate above; cauline leaves 2 or 3; cymes several to many flowered, the flowers showy; sepals 3-5.4 mm long, oblong- lanceolate, acuminate; petals yellow, 5-6.5 mm long; achenes several, 1.3-1.8 mm long; styles subapical, flliform, 1.8-2.1 mm long. Two moderately distinctive varieties are represented: Leaflets 11-15, uniformly long-strigose, not bicolored, with 5-19 teeth, often small, curled, and folded, without tomentum P. crinita var. crinita Leaflets 7-12, long-strigose below, tomentose above, somewhat bicolored, with 5-11 teeth, sometimes larger and flat; tomentose on stems, leaves, and petioles P. crinita var. lemmonii Var. crinita. {P. vallicoh Greene). Sage- brush, mountain brush, pinyon-juniper, pon- derosa pine, and aspen communities at 1890 to 2200 m in Wayne and Garfield counties; Arizona, New Mexico, and Colorado. The va- rieties are connected by intermediates; 10 (iii). Var. lemmonii (Wats.) Kearney & Peebles [P. lemmonii (Wats.) Greene; Ivesia lemmonii Wats.]. Mountain brush, pinyon-juniper, pon- 26 Great Basin Naturalist Vol. 42, No. 1 derosa pine, and aspen communities at 1900 to 2590 m in Garfield, Iron, Kane, Piute, and Wayne covmties; Arizona, Nevada, and New Mexico. This variety is intermediate between var. crinita and P. hippiana, but is distin- guished from P. hippiana by the narrow, few- toothed leaflets, strigose-glandular pu- bescence, and openly branched inflorescence with smaller flowers. To these crinita charac- teristics are combined the hippiana character of tomentum and bicolored leaves. Some few specimens are apparently exactly inter- mediate between var. lernmonii and P. hip- piana, and possibly represent products of hybridization; 20 (v). Potentilla diversifolia Lehm. Wedge-leaf Cinquefoil. Perennial herbs, the stems ascending to erect, 0.6-3.2 dm tall; leaves mainly basal, palmately to less commonly pinnately (3) 5- to 7-foliolate, the terminal leaflet 1.2-4 cm long, 0.3-1.8 cm wide, obo- vate to oblanceolate, toothed mainly above the middle, not bicolored, green on both sides, strigose to pilose on both sides, seldom tomentose below; cauline leaves 1-3; cymes several flowered, the flowers showy; sepals 5-6.5 mm long, triangular to triangular- attenuate; bracteoles 2.1-4.5 mm long, ob- long to lanceolate; petals yellow, 5.6-9.5 mm long; achenes numerous, 1.5-1.9 mm long; style subapical, 1.9-2.6 mm long, smooth or glandular, not basally thickened. Two varie- ties are present: 1. Leaflets divided 70-90 percent to midrib, into narrowly oblong segments; plants 15-20 cm tall, of the Uinta Mts P. diversifolia var. perdissecta Leaflets divided 40-70 percent to midrib, or usually merely toothed; plants 15-30 cm tall, widespread in mountains P. diversifolia var. diversifolia Var, diversifolia. {P. glmicophylla Lehm.; P. concinniformis Rydb.). Dry to wet mead- ows, lake margins, stream banks, forest mar- gins, alpine tundra, and rocky ridges at 2745 to 3500 m in Beaver, Cache, Daggett, Duch- esne, Garfield, Grand, Iron, Juab, Piute, Salt Lake, San Juan, Sevier, Summit, Tooele, Uin- tah, Utah, Wasatch, Washington, and Wayne counties; Alaska and Yukon, south to Califor- nia, Arizona, New Mexico, and South Da- kota. This is a highly variable taxon. The fol- lowing variety is the most pronounced of regional ecotypes, but is weak and difficult to distinguish in all cases; 50 (xv). Var, perdissecta (Rydb,) C, L, Hitchc. {P. perdissecta Rydb.). Rare in Utah, at 3100 m in the Uinta Mts., Summit Co.; Wyoming, Idaho, and Montana, and less commonly north in Canada; 38 (0). Potentilla effusa Dougl. ex Lehm, {P. colo- radensis Rydb.) Perennial herbs, multicipital from a branched caudex; stems ascending to erect, 2-4 dm tall; basal leaves 3.5-13 cm long, pinnately compound with 5-11 leaflets, the terminal one 2.0-4.0 cm long, 0.5-1 cm wide, oblanceolate to obovate, serrate or toothed about one half of way to midrib, the teeth only above the middle of the leaflet, sometimes apically few-toothed, white or grayish-tomentose above and below, never strongly bicolored; flowers several to many; sepals 3.5-4.7 mm long, lance-ovate, acumi- nate, conspicuously tomentose especially at anthesis; bracteoles 1.0-2.8 mm long, usually shorter than the calyx-lobes and darker in color; petals yellow, 4.0-6.5 mm long; ach- enes numerous; styles subapical, 1.9-2.8 mm long, filiform. Rare, rocky slopes and shelves of cliffs, north slope of the Uinta Mts. at 2740 m. Summit and Daggett counties; Colorado to central Alberta, almost always on the east- ern slope of the Continental Divide; 3 (i). Potentilla fissa Nutt, {Drymocallis fissa (Nutt.) Rydb.) Perennial glandular-pubescent herbs, low, 2-3 dm tall, from a caudex; basal leaves conspicuously veiny, dark green, pin- nately compound with 7-13 leaflets, the ter- minal one 1-5 cm long and 1-3 cm wide, broadly ovate to orbicular, deeply incised and doubly serrate, glandular-puberulent to glabrous on both surfaces; flowers several to many; sepals 5-10 mm long, long-acuminate, longer in fruit; bracteoles 4-6 mm long, nar- rowly lanceolate; petals yellow, orbicular, concave, 6-9 mm long; styles basally at- tached, 0.8-1.0 mm long, deltoid. Reported from Utah by Rydberg (1908, 1922); one specimen from Utah Co. at 2070 m.; south- western Colorado, southwestern Wyoming and central Idaho, north to Alberta and South Dakota, south to New Mexico. March 1982 Welsh: Utah Flora, Rosaceae 27 Potentilla fruticosa L. Shrubby Cinquefoil; Yellow Rose; Tundra Rose. [Fragaria fruti- cosa (L.) Crantz; Dasiphora riparia Raf.; D. fmticosa (L.) Rydb.; P. florihunda Pursh; PentophyUoides florihunda (Pursh) A. Love]. Shrubs to 1 m tall or more; bark shreddy; leaves 1-5 cm long, pinnately 3- to 7- foliolate, the terminal leaflet 0.5-2.5 cm long, 0.2-1 cm broad, oblong to elliptic, en- tire, green and sparsely hairy to glabrate above, grayish and silvery hairy below, some- what revolute; flowers 1 to several, con- spicuous; sepals 3.5-9 mm long, ovate-lan- ceolate; bracteoles 4-13 mm long, lanceolate to elliptic; petals yellow, 6-14 mm long, roimded; receptacle hairy; achenes 1.5-2 mm long, white-hairy. Meadows, sagebrush, as- pen, lodgepole, ponderosa pine, and spruce- fir communities often on floodplains or stream banks at 1700 to 3500 m in Carbon, Daggett, Duchesne, Garfield, Grand, Iron, Kane, Piute, San Juan, Salt Lake, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, and Wayne counties; Alaska east to New- formdland, south to California, New Mexico, Iowa, and New Jersey; Eurasia. This hand- some plant is known in cultivation in Utah, Summit, and Salt Lake counties at elevations below 1900 m, and should be grown widely in the state; 62 (ix). Potentilla glandulosa Lindl. Glandular Cinquefoil. [DnjmocaUis glandulosa (Lindl.) Rydb.] Perennial glandular-pubescent herbs, 0.8-6 (7) dm tall, from a caudex; basal leaves 3-22 cm long, pinnately compound, with 5-9 leaflets, the terminal one 0.7-6.8 cm long, 0.6-4.2 cm wide, obovate to elliptic, doubly dentate or lobed, green and variously pu- bescent or glandular on both surfaces; flowers several to many, showy or inconspicuous; sepals 5.5-9.3 mm long, lance-ovate, often acuminate, longer in fruit; bracteoles 4.5-7.2 mm long, oblong to narrowly lanceolate; pet- als mainly yellow, mostly 4-7.5 mm long; re- ceptacle sparsely hairy; achenes numerous, 1-1.2 mm long; styles from below the middle, 0.8-1 mm long. The glandular cin- quefoil is widespread and common in much of Utah, where it consists of a series of inter- grading populations varying by degree from each other and only arbitrarily separable from P. argiita (q.v.), with which it probably should be combined. In that case, the name would be P. arguta, because that epithet has priority. The complex was summarized by Keck (Carnegie Institution Washington 520: 26-124. 1940). He recognized four morpho- logical phases from Utah (i.e., the subspecies arizonica, micropetala, glabrata, and pseu- dorupestris). Examination of a fairly large series of specimens from Utah demonstrates that all, except that designated as micropetala are connected by a series of intermediates, and might best be regarded as belonging to a single polymorphic and highly plastic var. in- termedia. The following key will serve to dif- ferentiate most specimens. 1. Petals much shorter than the sepals, 4-5 mm long, 2-4 mm broad P. glandulosa var. micropetala - Petals shorter than to slightly exceeding the sepals, mainly 5-7.5 mm long and about as broad P- glandulosa var. intermedia Var. intermedia (Rydb.) C. L. Hitchc. [DnjmocaUis pseudorupestris var. intermedia Rydb.; D. glabrata Rydb.; P. glandulosa ssp. glabrata (Rydb.) Keck; P. glandulosa var. glutinosa f. glabrata (Rydb.) T. Wolf; D. arizonica Rydb.; P. glandulosa ssp. arizonica (Rydb.) Keck; P. macdougalii Tidestr.; P. pseudorupestris Rydb.; D. pseudorupestris (Rydb.) Keck]. Mountain brush, ponderosa pine, lodgepole pine, aspen, and spruce-fir commimities, often in meadows at 1890 to 3200 m in Beaver, Box Elder, Cache, Dag- gett, Duchesne, Garfield, Piute, Rich, Salt Lake, Sanpete, Sevier, Summit, Uintah, Utah, Wasatch, Washington, and Weber counties; British Columbia and Alberta south to Ore- gon, Arizona, and Wyoming; 33 (iv). Var. micropetala (Rydb.) Welsh & John- ston, comb. nov. [based on: Drymocallis mi- cropetala Rydberg, North Amer. Flora 22(4): 375. 1908. Type from Salt Lake Co.; P. glandulosa ssp. micropetala (Rydb.) Keck]. Sagebrush, mountain brush, upwards to al- pine meadows at 1430 to 3050 m in Salt Lake, Sanpete, Sevier, and Weber counties; Idaho and Wyoming. This small-petaled 28 Great Basin Naturalist Vol. 42, No. 1 phase resembles P. norvegica in flower size and in general conformation, but differs from inter alia in pistil features and leaflet num- ber; 4 (i). Potentilla gracilis Dougl. ex Hook. Slen- der Cinquefoil. Perennial, variably pubescent herbs from a caudex; stems ascending to erect, 0.4-6 (8) dm tall; basal leaves 3-30 cm long, or more, palmately compound, with 5-9 leaflets, the terminal one 1.3-10.7 cm long, 0.4-3.7 cm wide, obovate to oblanceo- late, crenate, serrate, or toothed to dissected, commonly slightly bicolored, green on both sides; flowers several to numerous, showy; sepals 3.5-9.5 mm long, lanceolate to lance- ovate, acute to attenuate-acuminate; brac- teoles 2.4-8.5 mm long, lance-oblong; petals yellow, 5.6-8 (10) mm long; achenes numer- ous, 1.3-1.6 mm long; styles subapical, 2-2.3 mm long, filiform or thickened to about half the length, tapered to stigma. Several inter- grading phases are recognizable at varietal level in this most common and widespread of our cinquefoil species. Variants tend to repre- sent recombinant types of several recurrent features: glands on calyx teeth, tomentum on lower leaflet surface, and depth of incision of leaflet margin. There are two varieties sepa- rable as follows: 1. Calyx lobes and often the leaflets glandular and stiffly pilose; plants often drying brownish p. gracilis var. brunnescens Calyx lobes and leaflets without glandular hairs; plants usually green P. gracilis var. glahrata Var. brunnescens (Rydb.) C. L. Hitchc. (P. brunnescens Rydb.). Meadows, mountain brush, aspen, spruce-flr, and alpine tundra at 2300 to 3180 m in Salt Lake, Sanpete, Sevier, Summit, and Utah counties; Washington and Montana south to Nevada and Wyoming. This variety is freely transitional to P. pul- cherrima and to P. diver sifolia, where they occur together; 9 (0). Var. glahrata (Lehm.) C. L. Hitchc. [P. nuttallii var. glabrata Lehm.; P. glahrata (Lehm.) Rydb.; P. chrysantha Lehm. in Hook., not Trev.; P. gracilis var. chrysantha (Lehm.) Rydb.; P. rigida Nutt.; P. gracilis var. rigida (Nutt.) Wats.; P. gracilis var. nuttallii Sheld.; P. gracilis ssp. nuttallii (Sheld.) Keck; P. hlaschkeana Turcz. ex Lehm.; P. gracilis var. hlaschkeana (Turcz.) Jeps.; P. viridescens Rydb.; P. gracilis var. viridescens (Rydb.) T. Wolf; P. glomerata A. Nels.; P. hlaschkeana var. glomerata (A. Nels.) T. Wolf; P. jucunda A. Nels.; P. diversifolia var. jucunda (A. Nels.) T. Wolf; P. grosseserrata Rydb.; P. rec- tiformis Rydb.; P. dichroa Rydb.; P. permollis Rydb.; P. gracilis var. permollis (Rydb.) C. L. Hitchc.]. Mountain brush, sagebrush, aspen, and spruce-fir communities at 1675 to 2745 m in Daggett, Piute, Sanpete, Sevier, and Weber counties; British Columbia and Al- berta south to California, New Mexico, and Nebraska. Included in our limited materials are those specimens with spreading hairs on petioles and stems, passing as var. permollis (Rydb.) C. L. Hitchc. They seem to be transi- tional completely with var. glabrata; 7 (i). Potentilla hippiana Lehm. Hipp Cinque- foil; Woolly Cinquefoil. [P. leucophylla Torr., not Pallas; P. pensylvanica var. hippiana (Lehm.) T. & C; Pentaphyllum hippianum (Lehm.) Lunnell; P. effusa var. filicaulis Nutt. in T. & C; P. filicaulis (Nutt.) Rydb.; P diffusa Gray; P. hippiana var. diffusa (Gray) Lehm.; P. hippiana var. propinqua Rydb.; P. propinqua (Rydb.) Rydb.; P. argyrea Rydb.; P. hippiana var. argyrea (Rydb.) B. Boi.]. Per- ennial, variably pubescent herbs from a cau- dex, the stems ascending to erect, 1.1-4.8 (5.5) dm tall; basal leaves 2.5-19 cm long or more, pinnately compound with 7-11 leaf- lets, the terminal one 0.9-4.7 cm long, 0.4-1.9 cm wide, oblanceolate to oblong or elliptic, serrate or toothed less than halfway to midrib, the teeth from below the middle, grayish-tomentose to pilose on one or both surfaces; flowers several to numerous, showy; sepals 4.3-6.5 mm long, lance-ovate, acute to acuminate; bracteoles 2.4-5.5 mm long, lance-oblong; petals yellow, 6-7.7 (9.5) mm long; achenes several to numerous, 1.5-1.9 mm long; styles subapical, 1.8-2.3 mm long. Meadow, aspen, spruce-fir, and alpine tundra communities at 2250 to 3450 m in Beaver, March 1982 Welsh: Utah Flora, Rosaceae 29 Daggett, Duchesne, Garfield, Grand, Iron, Piute, San Juan, Sevier, Summit, Uintah (?), Washington, and Wayne counties; British Columbia east to Michigan and south to New Mexico, Arizona, and Nebraska. Apparent in- termediates are known between the Hipp cinquefoil and P. gracilis, and especially P. pulcherrima; 18 (1). Potentilla hookeriana Lehm. Hooker Cin- quefoil. Perennial herbs, the stems ascending, 1-2 dm tall, from a caudex; leaves 1-3 cm long, all basal, ternate-digitate, the terminal leaflet 1-3 cm long, 0.5-1.5 cm wide, ovate to obovate, shallowly few toothed, densely white-tomentose below and sparsely tomen- tose to puberulent above, bicolored; petiole sericeous-strigose and tomentose; flowers 3-15, small; sepals 4-8 mm long, lanceolate- acmninate; bracteoles 3-6 mm long, lanceo- late-acuminate, densely villous and tomen- tose; petals yellow, obcordate, 2-5 mm long; achenes numerous; style subterminal, 0.8-1 mm long, usually uniformly thickened. Al- pine grassland and tundra, 3350 to 3360 m in Summit, Duchesne, Grand, and Juab coun- ties; Greenland to Alaska and south to Wyoming and Colorado; 4 (ii). PotentilUi multisecta (Wats.) Rydb. Dis- sected Cinquefoil. [P. diversifolia var. multi- secta Wats.]. Perennial herbs; stems decum- bent to ascending, 1-1.5 dm long, from a caudex; leaves (4) 5-8 (9) cm long, mainly basal, palmate to subpinnate, with 5-8 (12) leaflets, the terminal leaflet 1-4 cm long, and 1-2 cm wide, ovate to obovate, pinnately dis- sected into 3-8 long narrow segments, deeply divided, moderately to densely strigose, often grayish green; flowers 3-10, on recurved pedicels in fruit; sepals 3-6 mm long, strigose to strigulose; bracteoles 2-4 mm long, lan- ceolate-acuminate, strigose to strigulose; pet- als yellow, obcordate, 4-8 mm long; style subterminal, 2-1.3 mm long, filiform. Pinyon- juniper and sagebrush communities, rocky ridges of foothills at 2075 to 3000 m in Juab and Tooele counties; Nevada. Plants of open places are low, with the leaf rachis short, but shade foniis and those from rock crevices and ledges are large and lax. This entity had often been compared with P. diversifolia var. per- dissecta, but is distinctive in its strigose pu- bescence and recurved pedicels, and its habi- tat is at much lower elevations in drier sites. In habit, the plants resemble P. concinna, es- pecially its var. proxima, but differ in the much more dissected leaflets, absence of to- mentum, and recurved pedicels; 20 (iii). Potentilla nivea L. Perennial nonglandular herb, the stem decumbent to ascending, 0.7-2 dm long, from a caudex; stems and pe- tioles densely tomentose; basal leaves 2-9 cm long, trifoliolate, the terminal leaflet 0.8-2.7 cm long, 0.7-1.7 cm wide, obovate to elliptic or oblanceolate, coarsely toothed to near the base, green and silky-pubescent above, dense- ly snow white tomentose below, strongly bicolored; flowers 1-15 in an open in- florescence, showy; sepals 2.5-5 mm long, lanceolate; bracteoles 1.8-4.5 mm long, ob- long to lanceolate; petals yellow, 4-7 mm long; achenes several, 1-1.5 mm long; style subterminal, ca 1 mm long, uniformly thick- ened. Alpine tundra in the LaSal Mts., Grand and San Juan counties; Alaska east to Green- land, south in the Rocky Mountains to New Mexico and southeastern Utah; Eurasia; 8 (iii). This species is common and distinctive through the Rocky Mts., and probably should be expected in the Uinta Mts. as well. Potentilla norvegica L. Rough Cinquefoil. [Fragaria norvegica (L.) Crantz; P. mon- speliensis var. norvegica (L.) Farw.; P. mon- speliensis L.; Fragaria monspeliensis (L.) Crantz; P. norvegica ssp. monspeliensis (L.) Asch. & Graebn.; P. hirsuta (Michx.) Hylan- der]. Annual or biennial (short-lived per- ennial?) herbs, the stems erect, 0.7-5 dm tall or more, from a taproot, the stems and pe- tioles sparsely stiff-hairy; leaves mostly caul- ine, palmately (or subpinnately) compound with 3 (rarely 5) leaflets, the terminal one 1-8 cm long, 0.6-2.7 cm wide, obovate to ob- lanceolate, coarsely toothed to near the base or entire and cuneate in the lower part, green and sparsely stiff-hairy to glabrous above, paler and stiff-hairy beneath, espe- cially along the veins; flowers several to many, inconspicuous; sepals 4-6 mm long, ovate-lanceolate, enlarging in fruit; brac- teoles 3-6 mm long, oblong to elliptic or lan- ceolate; petals yellow or whitish, 2.5-3.6 mm long; achenes numerous, 0.8-1.1 mm long; style subterminal, 1.6-0.9 mm long. Flood- plains, wet meadows, lake shores and other moist sites at 1370 to 2930 m in Garfleld, Grand, Iron, Kane, Salt Lake, Sevier, Uintah, 30 Great Basin Naturalist Vol. 42, No. 1 and Utah counties; widely distributed in the northern hemisphere; 8 (i). Potentilla ovina Macoun Perennial herbs, from a caudex, the stems decumbent to as- cending, 1-3.5 (5) dm tall; leaves mainly bas- al, pinnately compound with (5) 9-18 leaf- lets, the terminal mostly 0.6-2.1 mm long, 0.4-1.1 cm wide, deeply pinnately dissected or apically few toothed, conspicuous; calyx 6-8 (10) mm long, lobes ovate-lanceolate to lance-attenuate; bracteoles 1.8-4 mm long, oblong to lance-oblong; petals yellow, 5-6.4 mm long; achenes many, 1.1-1.7 mm long; style subterminal, 1.8-2.7 mm long, filiform. There are two fairly distinct varieties in Utah: Leaflets densely to uniformly sericeous, grayish green or gray, often with a lower layer of sparse tomentum, 5-10 mm long, often with more than 6 teeth; leaf rachis 2-6 cm long P. ovina var.'ouina Leaflets glabrous to sparsely sericeous-strigose, usually green, never tomentose, 10-20 mm long, often with 3-5 teeth; leaf rachis 3-12 cm long P. ovina var. decurrens Var. decurrens (Wat.) Welsh & Johnston, comb. nov. [based on: Potentilla dissecta var. decurrens Wats., Proc. Amer. Acad. 8: 557, 565. 1873; P. nelsoniana Rydb.]. Meadows and rocky ridges, ponderosa pine, lodgepole, spruce-fir, aspen, and (less commonly) alpine tundra communities at 2200-3500 m in Bea- ver, Cache, Daggett, Duchesne, Garfield, Grand, Piute, San Juan, Sanpete, Summit, Tooele, Uintah, and Wayne counties; south- em Colorado west to central Nevada, north to southern Wyoming, more sparsely north- ward to Alberta. This is a characteristic form of montane meadows in the Uinta Mts., up to the alpine there and elsewhere. Var. ovina [P. diversifolia var. pinnatisecta Wats.; P. wyorningensis A. Nels.; P. mon- idensis A. Nels.]. Meadows and rocky ridges, openings in spruce-fir and alpine tundra com- munities, 2800-3900 m in Cache, Davis, Duchesne, Grand, Juab, Piute, Salt Lake, San Juan, Sanpete, Utah, and Summit counties; southwestern Alberta south to northeastern Oregon, central Utah, and southern Wyom- ing. This taxon has been confused with P. plattensis, but its tomentulose leaflets and erect-ascending pedicels in fruit appear to be diagnostic. Hybrids between P. ovina (usually var. decurrens) and P. pulcherrima are fre- quently encountered in Utah (Box Elder, Cache, Piute, Daggett, and Summit counties); leaves are pinnate or subpalmate with 7-15 leaflets that are usually bicolored and moder- ately tomentose beneath, and stems are ascending, contrasting with the usually de- cumbent ovina and the erect-ascending pulcherrima. Potentilla paradoxa Nutt. in T. & G. Con- trary Cinquefoil. [P. supina Michx., not L.; TridophyUiim paradoxum (Michx.) Greene; P. supina var. nicolletii Wats.] Annual, biennial or short-lived perennial herbs, the stems de- cumbent to ascending or erect, 0.8-9 dm tall, sparsely to moderately villous; leaves mainly cauline, all pinnately compound with 5-11 leaflets, the terminal one 1.2-4.5 cm long, 0.4-2.5 cm wide, often dissected into 3 main confluent lobes, obovate to oblanceolate, toothed to below the middle, cuneate to the base, green on both sides, strigose to strigu- lose; flowers few to many, not especially showy; sepals 3-5.2 mm long, ovate, shorter than the bracteoles, enlarging in fruit; brac- teoles 4-6.5 mm long, lanceolate; petals yel- low or whitish, 3.2-4.3 mm long; achenes nu- merous, 0.8-1 mm long, laterally enlarged along ventral suture; style subterminal, 0.5-0.7 mm long. Beaches, marshes, and lake shores at 1350 to 1650 m in Salt Lake and Utah counties; widespread in North America; Asia; 6 (0). Potentilla pectinisecta Rydb. [P. Candida Rydb.; P. pecten Rydb.] Perennial herbs, 3-4 dm tall; stems stout and erect to ascending from a caudex, silky-strigose; leaves mostly basal, strictly palmate with 5-9 leaflets, the leaflets oblanceolate, silky-strigose on both sides, sometimes also sparsely tomentose March 1982 Welsh: Utah Flora, Rosaceae 31 beneath, deeply pectinately divided in ob- long-linear segments; inflorescence glomer- ately many-flowered with relatively thick pedicels; flowers large and showy; calyx silky-sericeous, 6-9 mm long, with lanceo- late-acmninate lobes; petals yellow, obcor- date-emarginate, TVs mm long; styles uni- formly thickened about one-half the length then filiform above, 1.8-3.2 mm long. Rare, at 2150 to 2750 m in Tooele and Salt Lake counties; Wyoming (?). Potentilla pensylvanica L. Perennial, glandular to nonglandular herbs from a cau- dex, the stems ascending to erect, 0.5-4.5 dm tall or more; leaves mostly basal, 2-18 (25) cm long, erect, pinnately compound with 5-17 leaflets, the terminal one 0.9-4.5 (6) cm long and 0.4-1.2 cm wide, elliptic to oblong or oblanceolate, coarsely toothed to narrowly lobed, the sinuses extending more than half way to the midrib, conspicuously revolute margined, green and somewhat hairy above, white or more commonly greenish or yellow- ish tomentose below; flowers few to many in a glomerate inflorescence, showy; sepals 3.5-7.5 mm long, ovate-lanceolate; brac- teoles 3-6 mm long, narrowly lanceolate; petals yellow, 5-8 mm long; receptacle glabrous; achenes numerous, 0.9-1.2 mm long; style subterminal, 0.9-1.0 mm long, coniform, with a conspicuously thickened and often pappillose base. This is a highly variable species throughout its large range in North America; the taxonomic problems in this species are mirrored in the whole section (sect. Multifidae), to which P. mbricaulis and P. multifida belong. These are misunderstood and complex species of the western and east- ern hemispheres, respectively. There is seem- ingly only one entity in Utah that merits sep- aration from typical P. pensylvanica: 1. Leaflets 5-7, subdigitate (10-30 percent of the rachis occupied with leaflets); pubescence silvery or yellowish white, often of long entangled hairs densely matted; plants alpine. La Sal and Tushar Mts P. pensylvanica var. paucijuga - Leaflets 7-17, often pinnate (30-60 percent of rachis occupied); pubescence olive greenish, dull, not silvery, usually composed of short curly hairs sparsely matted; plants widespread, rarely alpine in Uinta Mts P. pensylvanica var. pensylvanica Var. paucijuga (Rydb.) Welsh & Johnston, comb. nov. [based on: Potentilla paucijuga Rydb. N. Amer. Fl 22(4): 348. 1908. Type from La Sal Mts., Putjius 251 in August 1899 (US! photo NY!)]. Grassy tundra, alpine com- munities, at 3800 m in Grand, Piute, and San Juan counties; endemic to Utah. In the LaSal Mts. this form may hybridize with P. nivea; the two species are usually distinguished by the glomerate inflorescence with glandular pubescence, revolute-margined leaflets, and pinnate leaves of var. paucijuga. Another va- riety of this species occurs in similar situa- tions in California (var. ovium Jeps., not P. pseudosericea Rydb.), and it is possible that other alpine ecotypes of P. pensylvanica de- serve recognition in the Great Basin; 11 (ii). Var. pensylvanica [P. pseudosericea Rydb.; P. arachnoidea (Lehm.) Dougl. ec Rydb.] Sagebrush, sagebrush-grass, and meadow commimities at 2200-3450 m in Beaver, Car- bon, Duchesne, Emery, Garfield, Grand, Iron, Sevier, Wayne, and Wasatch counties; Hudson Bay to Alaska, south to northern Mexico, Texas, and Arizona. 38 (v). Potentilla plattensis Nutt. [Ivesia pinnati- fida Wats.; P. arizonica Greene; P. plattensis var. pedicellata A. Nels.]. Perennial herbs with stems 0.5-2.5 dm long, decumbent or creeping through meadow herbs and grasses; leaves basal and cauline, 3-11 cm long, pin- nately compound with 11-23 leaflets, these verticillate or subverticillate; terminal and lower leaflets all pinnately toothed with 5-8 teeth cutting 70-90 percent to midrib, glabrous to sparsely strigose, green to grayish green on both surfaces; stems with 3-15 flow- ers on recurved pedicels in fruit; calyx (5) 6-8 mm long, strigose or puberulent; achenes numerous; styles subterminal, 1.5-3.0 mm long, filiform. Wet meadows, bogs, and val- ley bottoms, rare in Utah, 1820-2700 m in Cache, Kane, and Sevier counties; Alberta and Manitoba south through Wyoming and 32 Great Basin Naturalist Vol. 42, No. 1 Colorado to central Arizona and New Mexi- co. This species has been confused with P. ovina in the past, but the habitats are usually sharply distinct, and the combination of trail- ing stems, pinnately dissected leaflets, sub- glabrous pubescence, and recurved pedicels are diagnostic for P. plattensis. Sometimes there are two forms at the same site: a lax trailing form in moist portions of a meadow, and a more compact form with prostrate stems on better-drained soil in the same meadow. Potentilla pukherrima Lehm. [P. filipes Rydb.; P. wardii Greene; type from Thousand Lake Mtn., Wayne Co.]. Perennial herbs, 3-8 dm tall, erect to ascending from a caudex; leaves mostly basal on long petioles, 8-25 cm long, palmate or subpinnate with 7 (9) leaf- lets, the terminal leaflet 1.5-8 cm long and 1-3.5 cm wide, oblanceolate or spatulate, with 10-15 coarse teeth cutting less than one-half, sericeous or strigose above, densely tomentose below, strongly bicolored; stem with 10-40 flowers that are large and showy; calyx large, 8-12 mm high, with acuminate lobes, sericeous-strigose and sometimes glandular in addition; petals yellow, 7-14 mm long; style filiform or slightly thickened at base, 1.6-3.0 mm long. Meadows and rocky slopes, pinyon-juniper, ponderosa pine, mountain shrub, and aspen communities at 2200 to 3380 m in Beaver, Box Elder, Car- bon, Daggett, Garfield, Grand, Iron, Piute, San Juan, Duchesne, Emery, Salt Lake, San- pete, Sevier, Summit, Uintah, Utah, Wasatch, and Wayne counties; British Columbia east to Quebec, south to New Mexico, Arizona, and Nevada; 66 (xiii). This species has often been compared with P. gracilis, but seems distinctive in its bicolored leaves, style form, oblanceolate leaflets, glandular pubescence, and longer petioles. It has also been confused with P. concinna, but contrasts with the stri- gose pubescence, prostrate habit, few- flowered inflorescence, and the simply acute calyx lobes and bracteoles of that species. Hybrid swarms are fairly frequent between P. pulcherrima and P. hippiana (Garfield, Grand, Piute, San Juan, Sevier, Summit, and Wayne counties), in which any combination of the distinguishing characters may be found. Potentilla rivalis Nutt. [P. millegrana Eng- elm. ex Lehm; P. leucocarpa Rydb. in Britt. & Br.] Annual or biennial herbs from a taproot, the stems 2-6 dm tall, spreading to suberect, pubescent with fine villous or villous-tomen- tose hairs; leaves mainly cauline, palmately to subpinnately compound with 3-5 leaflets, the terminal 1-4 cm long, 0.4-1.3 cm wide, obovate to oblong to oblanceolate, coarsely toothed (at least above the middle), green and minutely villous on both sides; flowers numerous, inconspicuous; sepals 2.3-4.2 mm long, ovate; bracteoles 3.3-4 mm long, lan- ceolate; petals yellow, 1.5-1.9 mm long, not enlarged laterally; style subterminal, 0.6-0.8 mm long, fusiform. Reservoir shore at 2300 m in Carbon Co., and to be expected elsewhere; widespread in North America; 1 (i). Potentilla rubricaulis Lehm. [P. saximon- tana Rydb; P. pedersenii Rydb.; P. furcata Porsild]. Perennial herbs, stems ascending from a caudex, leaves mostly basal, eglandu- lar, palmate to subpinnate with 5-7 leaflets, the terminal one 1.0-2.5 cm long and 0.5-1.0 cm wide, few toothed with narrow-crenate or serrate teeth, usually deeply divided 50-90 percent, often revolute margined, densely grayish-tomentose below, sparsely tomentose to puberulent above, strongly bicolored, of- ten pilose as well below with tufts of hair at the tips of ultimate segments, the petioles pi- lose-sericeous with subsidiary tomentum; cymes 1- to 10-flowered, the flowers showy but small; sepals 4-6 mm long; bracteoles 3-6 mm long; style subapical, 0.9-1.1 mm long, conic, usually conspicuously thickened and papillose below. Alpine tundra meadows and rocky ridges, 3600 to 4000 m in Du- chesne, Grand, Piute, San Juan, and Summit counties; Greenland to Alaska, south to Brit- ish Columbia, sparsely and rarely through Montana into Wyoming, and Colorado. This name is very often misapplied, but here it is taken for a strictly alpine species with cone- shaped, thickened style about one mm long, more or less digitate leaves with 5-7 leaflets, and petioles with long straight hair in addi- tion to tomentum. It has affinities with sect. Multijugae (see P. pensylvanica), but may be related to P. nivea and its relatives as well, with which it may hybridize; 5 (0). March 1982 Welsh: Utah Flora, Rosaceae 33 Prunus L. petals 5, white, pink, rose, or red; stamens numerous; pistil 1, free from the usually de- Shrubs or small trees with unarmed ciduous hypanthium; style 1, elongate, with branches; leaves alternate, simple, entire to capitate stigma; fruit a drupe. Note: The serrate or serrulate, rarely lobed, commonly genus is represented by only two mdigenous with glands on petioles or blade bases; sti- species in Utah, P. fasciculata and P. vir- pules lance-attenuate to linear, caducous; giniana. However, the numerous cultivated flowers perfect, regular, solitary, in umbel- species persist following cultivation, and late clusters, or in racemes; sepals 5, borne many escape. They are treated herein for atop a cup-shaped to turbinate hypanthium; those reasons. 1. Leaves entire or serrate only near the apex, borne in fascicles, less than 2 cm long and 0.5 cm wide; shrubs with divaricate branches, of low elevations, indigenous in Millard, Beaver, and Washington counties P. fasciculata - Leaves various, usually larger; shrubs to trees of moderate size, variously distributed, but not of desert shrublands 2 2(1). Flowers borne in pedunculate, elongate, or corymbose clusters 3 - Flowers borne singly or in sessile umbellate clusters from winter buds 7 3(2). Flowers borne in corymbose racemes, the pedicels subtended by persistent bracts - Flowers borne in elongate racemes, the pedicels without persistent bracts 5 4(3). Flowers to 2 cm wide or more, the petals often pink or double or both; bracts 3-8 mm long, cuneate, the truncate apex fringed-serrulate; flowering cherry .... P. serrulata - Flowers ca 0.8 cm wide, the petals white, single; bracts ca 1 mm long, ovate, serrulate; escaping (rootstock) P- mahaleb 5(3) Leaves leathery, thickened, more or less evergreen, entire; plants cultivated shrubs P hurocerasus - Leaves not especially thickened, serrate to serrulate; shrubs or small trees, cultivated or not 6(5). Hypanthium pubescent within; racemes pendulous or spreading in anthesis; cultivated ornamental ^- P^^"^ - Hypanthium glabrous within; racemes erect or ascending in anthesis; indigenous or less commonly cultivated P- virginiana 7(2). Leaves cordate-ovate to broadly ovate; cultivated apricot, rarely escaping P. armeniaca - Leaves lanceolate to oblong, or less commonly oblanceolate to broadly elliptic; cultivated ornamental and fruit trees and shrubs " 8(7). Plants shrublike, mainly 2 m tall or lower ^ - Plants shrub- or treelike, mainly 2-5 m tall or more H 9(8). Leaves glabrous, less than 2 cm broad; flowers commonly 2-4 per bud P. besseyi - Leaves hairy ,at least along veins beneath; flowers solitary (rarely 2) per bud 10 10(9). Leaves long-villous beneatii, sparingly villosulose above, broadly elliptic to oblong; flowers single; cultivated for ornament and for fruit P. tomentosa - Leaves sparingly villous on both sides; flowers mostly double; cultivated ornamental ....:.' ^- ^"^^« 34 Great Basin Naturalist Vol. 42, No. 1 11(8). Flowers solitary, or sometimes 2 or 3 per bud 12 — Flowers 3 or more per bud 15 12(11). Axillary buds on fruiting branchlets borne in 3s, the lateral ones producing flowers 13 — Axillary buds on fruiting branchlets borne singly 14 13(12). Leaves pubescent beneath; pedicels pubescent; cultivated plum P. domestica — Leaves glabrous beneath, except on midrib; pedicels glabrous; cultivated flowering plum P. ceracifera 14(12). Leaves sharply serrate; fruit not leathery and splitting, exposing the stone at maturity; cultivated peach, commonly escaping p. persica — Leaves crenate-serrate; fruit leathery, splitting and exposing stone at maturity; sparingly cultivated almond, not escaping p. dulcis 15(11). Plants fomiing shrubby thickets; flowers 14 mm across or less; fruit a plum P. americana — Plants treelike or small trees, not forming thickets; flowers 10-30 mm across or more; fruit a cherry 16 16(15). Inner flowerbud scales reflexed or spreading; petals 8-14 mm long; sweet cherry P. avium — Inner flower bud scales erect; petals 6-9 mm long; tart cherry P. cerasus Prunus americana Marsh. American Plum; Pottawattami Plum. Shrubs, often forming thickets, rarely treelike, to 5 m tall; branch- lets sometimes thomlike, glabrous; leaves 2.5-7 cm long, 0.5-3 cm wide, elliptic to ovate or lanceolate, sharply serrate, long-at- tenuate apically, acute to obtuse basally, glabrous or pubescent along veins beneath; petioles usually glandless; flowers 1-4, in ses- sile or subsessile umbels, from lateral buds, appearing before the leaves, in ours mainly 14 mm broad or less; petals white, 5-7 mm long, 2.5-3 mm wide; sepals spreading, pu- berulent; hypanthium puberulent without and within; fruit a yellow to red plum. Culti- vated fruit plant, and probably indigenous in portions of Utah, but also introduced by pio- neers to all areas of the state. The plants spread underground and form thickets that persist. Specimens examined were from Duchesne, Uintah, Utah, and Wayne coun- ties; introduced from central United States; 5 (2). Prunus armeniaca L. Apricot. Small trees to 8 m tall or taller; branchlets green to brown, armed or imarmed; leaf blades mainly 1.5-7 cm long, 1-6 cm wide on mature branches, cordate-ovate to ovate, obtusely serrate, abruptly attenuate, obtuse to cordate basally, glabrous, or hairy along veins be- neath; petioles usually with glands; flowers solitary, appearing before leaves; petals white (rarely p hkish), 8-12 mm long, obcor- date to orbicular; sepals glandular, the hy- panthium glabrous except basally; pedicels villosulose; fruit pubescent, fleshy and edible. Apricot trees are grown in the lower eleva- tion portions of the state, where they escape and persist as waifs along canals, roadsides, and fence rows. Specimens examined are from Beaver, Carbon, Millard, Utah, and Wayne counties; introduced from China; 8 (iii). Prunus avium L. Sweet Cherry. Trees to 8 m tall or taller; branchlets soon brown, unarmed; leaf blades mainly 3-10 cm long, 2.5-8 cm wide, broadly oblanceolate to obo- vate or elliptic, obtusely serrate to doubly serrate, abruptly short-attenuate, obtuse to rounded basally, glabrous or long-hairy espe cially along veins beneath; petioles com- monly with glands; flowers 2-4 per bud, ap- pearing with early leaves; petals white (rarely pink), 8-14 mm long, obcordate to or- bicular; sepals, hypanthium, and pedicels glabrous; fruit glabrous, red to almost black, edible. The Bing, Lambert, Royal Ann, and other cultivars of the sweet cherry are grown March 1982 Welsh: Utah Flora, Rosaceae 35 commercially and in garden orchards at low elevations in Utah, where frost is not prohibi- tive. Trees escape from cultivation along canals and in riverbottoms. Many are grown as grafted stock; some on rootstocks of other Pninus species, especially on P. mahaleb (q.v.). Specimens examined are from Carbon, Utah, and Washington counties; introduced from Eurasia; 4 (0). Prunus besseyi Bailey Western Sand Cherry. Shrubs to 1.3 m tall; branchlets soon brownish, not thornlike; leaf blades mainly 1.5-5 cm long, 0.5-1.8 cm wide, elliptic to oblanceolate, obtusely serrate, acute to cuspi- date apically, cuneate basally; glabrous; pe- tioles commonly lacking glands; flowers com- monly 2-4 per bud, appearing with early leaves; petals white, 4-6 mm long, 2.5-3.5 mm wide, elliptic; sepals, petals, and hypan- thium glabrous; fruit glabrous, black, com- monly astringent. The western sand cherry is used as a dwarfing rootstock for peaches, cherries, and other Primus species. The plants persist and escape in orchard areas of the state; erosion control and wildlife plant- ings are probable; specimens were examined from Utah and Wasatch counties; introduced from the Great Plains; 3 (i). Prunus cerasifera Ehrh. Cherry Plum; Flowering Plum. Small trees to 6 m tall, rarely taller; branchlets soon brown, not thomlike; leaf blades mainly 1.8-6.5 cm long, 1.2-4 cm wide, ovate to elliptic, serrate to doubly serrate, acute apically, acute to rounded basally, villous along the veins be- neath; petioles seldom with glands; flowers solitary, or less commonly 2 or 3 per bud; petals pink to violet or white, 6-9 mm long, 4.5-7 mm wide, suborbicular; sepals, hypan- thium, and pedicels glabrous, except along marginal insertion of filaments; fruit red to purple, edible. Cultivated ornamentals of streets, lawns, and gardens, persisting and es- caping in lower elevation portions of the state; specimens examined from Utah Co.; in- troduced from Asia; 8 (i). The hybrid be- tween P. cerasifera and P. mume Sieb. & Zucc, the Japanese apricot is also cultivated in Utah, under the name P. x blireana Andre. The hybrid has broad purple leaves. Prunus cerasus L. Sour Cherry; Pie Cherry. Small trees to 5 m tall, rarely more; branchlets soon brown, not thornlike; the leaf blades mainly 3-10 cm long, 1.2-5 cm wide, oblanceolate to obovate or elliptic, doubly serrate, abruptly acuminate, acute to rounded basally, glabrous except hairy along veins beneath; petioles bearing glands; flow- ers commonly 3 per bud (less commonly few- er); petals white, 7-9 mm long and about as broad; sepals crenate-serrate, glabrous; hy- panthium and pedicels glabrous; fruit red, soft, sour. This is the tart cherry of com- merce, widely cultivated in Utah and a favor- ite food of robins; specimens examined from Utah Co.; introduced from Eurasia. The spe- cies flowers later than does the sweet cherry, and fruit is harvested in mid- to late July, af- ter the sweet cherries have been harvested. The trees long persist; 1 (0). Prunus domestica L. Common or Eu- ropean Plum; Italian Prune. Small trees to 5 m tall, with grayish or ashy bark; branchlets pubescent when young, not especially thorn- like; leaf blades mainly 2-10 cm long and 1-6 cm wide, ovate to obovate, coarsely ser- rate, rough and thinly pubescent above, pu- bescent beneath; petioles commonly with glands; flowers solitary or sometimes 2 or 3 per bud; petals white, ca 10 mm long; sepals and hypanthium pubescent or glabrous; pedi- cels commonly pubescent; fruit commonly blue purple, glaucous. Included here are the yellow-fleshed plums known as Damson and Green-Gage, other plums, and the Italian prune. The species is widely cultivated in Utah, where it persists and less commonly es- capes. The red-fleshed plums, called Japanese or Satsuma, belong to P. salicina Lindl. and are cultivated in the area too, but less com- monly; specimens were examined from Bea- ver and Utah counties; introduced from Eu- rasia; 4 (i). Prunus dukis (Mill.) D.A. Webb. Almond. Small trees to 5 m tall or taller; branchlets pale, not thornlike; leaf blades 2-7 cm long, 0.7-2 cm long on mature branches, oblong- lanceolate, crenate-serrate, abruptly short- apiculate, glabrous; petioles usually bearing glands; flowers solitary, appearing before or with early leaves; petals pink; sepals villous at margin; fruit pubescent, splitting at matu- rity and exposing the stone. Sparingly culti- vated ornamental, botanical curiosity, and 36 Great Basin Naturalist Vol. 42, No. 1 "nut" tree, known from Utah and Washing- ton counties; persisting, but not escaping (?); introduced from the Old World; 1 (0). Prunus fasciculata (Torr.) Gray. Desert Peach. [Emplectocladus fasciculatus Torr.]. Low intricately branched shrub to 1.5 (2) m tall; branchlets pubescent, ashy or grayish, more or less thomlike; leaves mainly 0.5-2.5 cm long, 0.1-0.6 cm wide, cuneate-spatulate, entire or few toothed near the apex, apicu- late or cuspidate, sessile or nearly so; not bearing glands at leaf base, more or less pu- berulent on both sides; flowers mainly 1 per bud; petals cream to white, 3.5-5 mm long, spatulate to elliptic; sepals, hypanthium, and pedicels glabrous; fruit hairy, thin fleshed, in- edible. The desert peach occurs in mixed desert shrub and lower pinyon-juniper com- munities at 625-1770 m in Beaver, Millard, Iron (?) and Washington counties; Arizona, Nevada, and California; 27 (vii). Prunus laurocerasus L. Cherry-Laurel. Evergreen shrub to 2 m, rarely more; branch- lets pallid, glabrous, not at all thornlike; leaf blades mainly 3-10 (13) cm long, and 1-3 cm wide, entire or remotely crenate-serrate, el- liptic to oblong, attenuate to abruptly so, cu- neate to acute basally, leathery-thickened, glabrous; petioles lacking glands; racemes not leafy, commonly 3-10 cm long, many flow- ered; the pedicels 0.5-1.5 mm long, sub- tended by large caducous bracts; flowers ca 1 cm wide or less; petals white, 3-4 mm long, obovoid; sepals fringed, glabrous; hypan- thium and pedicel glabrous; fruit black, ined- ible. Cultivated ornamental; specimens from Davis, Utah, and Weber counties; introduced from Eurasia; 8 (0). Prunus mahaleb L. Mahaleb; St. Lucie Cherry. Small trees to 6 m tall or taller; branchlets pale brown, copiously pubescent, not thornlike; leaf blades mainly 2-6 cm long, 1.5-4.5 cm wide, oval to elliptic or ovate, finely crenulate, abruptly acute, acute to rounded basally, commonly glabrous on both sides; petioles sometimes with glands; racemes with 3-12 flowers, corymbose, the axis short, leafy bracted at base; petals white, 4-6 (8) mm long, oblong-oblanceolate; sepals, hypanthium, and pedicels glabrous; fruit black. The Mahaleb Cherry is used as root- stock for other cherry cultivars. It persists and escapes, becoming established along canals and in riverbottom forests in Utah Co.; introduced from Asia; 5(1). The bitter cherry P. emarginata (Dougl.) Walpers is reported for Utah in Vascular Plants of Pacific North- west 3: 160. 1961, but I have not seen any material of that species from Utah. It would key to P. mahaleb in the above key, but differs from inter alia in usually lacking leafy bracts on the peduncle, and in the rounded to obtuse or rarely acute leaves. In some bitter cherry specimens the stems are glabrous and the calyx is hairy. The petals are often pu- bescent on the dorsal surface, and stones are spindle shaped. Prunus padus L. European Bird Cherry; May Day Cherry. Small trees to 8 m tall, rarely taller; branchlets soon brown, glabrous or puberulent, not thornlike; leaf blades 1.5-20 cm long, 0.7-3.5 cm wide, elliptic to oblanceolate or obovate, serrate, abruptly acuminate, acute to truncate basally, glabrous except sometimes on veins beneath; petioles usually bearing glands; racemes with leafy peduncles, commonly 7-12 cm long, 15- to 25-flowered, the pedicels 5-17 mm long, subtended by caducous bracts; flowers 12-20 mm broad; petals white, 5-7.5 mm long, almost as broad; sepals fringed, glabrous; hypanthium and pedicels glabrous; fruit black, bitter, astringent. Cultivated or- namental of yards and other plantings; Utah Co.; introduced from Eurasia; 1 (0). Prunus persica (L.) Batsch. Peach. Small trees to 3 (4) m tall, seldom more; branchlets green to pallid, becoming ashy in age, glabrous, not thornlike; leaf blades mainly 3-15 cm long (or more), 0.5-5.5 cm wide, ob- long-lanceolate to lanceolate, serrate to cre- nate-serrate, attenuate, obtuse to acute ba- sally, glabrous; petioles usually bearing glands; flowers solitary, appearing before the leaves; petals pink, white, or red; sepals vil- lous at margin; fruit pubescent, fleshy at maturity, edible. Cultivated fruit and orna- mental trees at lower elevations in the state, widely escaping along roads; specimens seen from Carbon, Utah, and Salt Lake counties; introduced from China; 8 (i). Prunus serrulata Lindl. Flowering Cherry. Trees to 10 m tall or more; branchlets pallid, becoming brown, glabrous, not thornlike; leaf blades mainly 3-15 cm long, 1.2-8 cm wide. March 1982 Welsh: Utah Flora, Rosaceae 37 ovate to lance-ovate, abruptly long-acumi- nate, aristate-serrate, acute to obtuse basally; petioles usually bearing glands; racemes co- rymbose, 3- to 5-flowered, naked at the base, the pedicels 1.5-30 mm long, each subtended by a cuneate bract fringed at the truncate apex; petals 12-20 mm long, oval, white, rose, pink, often double; sepals glabrous, acuminate or toothed; hypanthium and pedi- cels glabrous; fruit small, black. This flower- ing cherry is gaining in popularity, especially in grafted or pendulous forms; specimens ex- amined from Utah Co.; introduced from Ja- pan; 4 (0). Prunus tomentosa Thunb. Bush Cherry. Shrubs to 2 m tall (commonly lower); branch- lets brown, copiously pubescent, not thorn- like; leaf blades mainly 2-6 cm long, 1.5-3 cm wide, obovate to elliptic, abruptly acumi- nate, doubly serrate and obscurely lobed (in some), puberulent above, long-villous be- neath; petioles pubescent, not bearing glands; flowers 1 (rarely 2) per bud, appearing before the leaves, sessile or subsessile; petals white or pink, 7-9 mm long, oval; sepals serrulate, pilose; hypanthium glabrous below; fruit red, sparinglv pubescent, sour, edible. Cultivated ornamental, escaping and persisting in Utah Co.; introduced from Asia; 4 (0). Prunus triloba Lindl. Flowering Almond. Shrubs to 2 m (rarely taller); branchlets brown, glabrous; leaf blades commonly 2-5 cm long, 1.5-4 cm broad, ovate to obovate, sharply doubly serrate, pubescent on both sides; flowers 1 or 2 per bud, short-pedicel- late; petals pink or white, 8-15 mm long, oval, commonly double; sepals serrulate, glabrous; hypanthium glabrous; pedicels pu- berulent; fruit red, seldom produced. Spar- ingly cultivated shrubs, in Salt Lake and Utah counties; introduced from China; 1 (0). Prunus virginiana L. Chokecherry. Shrubs or small trees to 8 m tall with ashy bark; branchlets brown, glabrous, not thornlike; leaf blades 2-10 cm long, 1.5-7 cm wide, el- liptic to oblong-ovate, serrate, abruptly acuminate, acute to rounded basally, glabrous or sometimes pubescent beneath; petioles usually bearing glands; racemes 4-20 cm long, the peduncles usually leafy, 2-8 cm long; flowers 10-20 mm wide, numerous, the pedicels 4-17 mm long, each subtended by a caducous bract; petals white, 4-6 mm long. suborbicular; sepals fringed, glabrous; hypan- thium and pedicels glabrous; fruit black when ripe, astringent, edible. Chokecherry fruit has been gathered since early days for making of jelly and syrup, and prior to that by indigenous peoples as a component of pemican. The species is known from all coun- ties in Utah. Our material has been assigned to var. melanocarpa (A. Nels.) Sarg. [Cerasus demissa var. melanocarpa A. Nels.; P. me- lanocarpa (A. Nels.) Rydb.; F. demissa var. melanocarpa (A. Nels.) A. Nels.]. Reports of P. demissa or of P. virginiana var. demissa (Nutt.) Torr. for Utah belong to var. melano- carpa. That variety is widely distributed in western North America; 158 (xxvi). PuRPUSiA Brandegee Plants perennial, caespitose, glandular herbs, arising from a caudex; leaves mainly basal, pinnately compound; flowers perfect, regular, borne in few-flowered cymes; hypan- thium campanulate to turbinate, usually lack- ing bractlets; sepals 5; petals 5, white to yel- lowish; stamens 5, opposite the sepals; pistils mostly 6-12, on a stalked receptacle; fruit of achenes. Purpusia saxosa Brandegee [P. arizonica Eastw.; P. osterhoutii A. Nels.; Potentilla os- terhoutii (A. Nels.) J. T. Howell]. Perennial glandular herbs from a caudex, the stems 0.5-2 dm tall; basal leaves pinnate with 5-11 leaflets, the leaflets 0.5-1.5 cm long, broad, deeply toothed or cleft; sepals 2.5-3 mm long, lance-ovate, acuminate; petals yellow or white, 3-4 mm long, oblanceolate, acumi- nate; achenes on a receptacle 1.5-2 mm long. This species is included in Utah on the basis of a collection reported by Meyer (1976) taken at Kolob Reservoir, Washington Co.; Arizona, Nevada, and California; 0 (0). PuRSHLA DC. ex Poir. Shrubs with unarmed branches; leaves al- ternate, simple, apically 3-toothed, usually glandular; stipules triangular-attenuate, per- sistent; flowers perfect, regular, solitary, on short lateral spur branchlets; sepals 5, borne atop a turbinate-fvmnelform persistent hy- panthium; petals 5, yellow; stamens ca 25; pistils 1 or 2, borne on a stipe at base of hy- panthium; fruit an achene. 38 1. Great Basin Naturalist Vol. 42, No. 1 Leaves with conspicuously punctate, depressed glands, glabrous above; plants of Washington Co P. glanduhsa Leaves lacking punctate depressed glands, puberulent above; plants widespread P. tridentata Purshia glandulosa Curran Shrubs, much branched, 1-2 (3) m tall; branchlets promi- nently glandular; leaves 3-10 mm long, 1- to 4-mm wide, cuneate, glabrous above, slightly tomentose beneath, the margins re volute; hy- panthium glabrous to tomentose, 2.5-5 mm long, funnelform; petals 4-8 mm long, spatu- late, creamy white to yellowish; achenes oblique, ca 2 cm long, including style, pu- berulent. Blackbrush, chaparral, and pinyon- juniper communities at 1065 to 1375 m in Washington Co.; Nevada, Arizona, and Cali- fornia; 2 (i). Purshia tridentata (Pursh) DC. Bitter- brush. Shrubs, much branched, to 2 m tall (rarely taller); branchlets brown, tomentu- lose; leaves mainly 4-20 mm long, 2-12 mm wide, cuneate, apically 3 (5) -toothed, tomen- tose but green above, grayish tomentose be- neath, the margins more or less re volute; hy- panthium tomentose and stipitate glandular; sepals mainly 4-6 mm long, ovate-oblong, entire; petals 5-9 mm long, oblong to obo- vate or spatulate, yellow; achenes obliquely ovoid, 1-2 cm long, the beak about one-third the length, puberulent. Bitterbnish is a plant of sagebrush, mountain brush, pinyon-juni- per, and ponderosa pine communities at 1220 to 2775 m in all counties in Utah; British Co- lumbia east to Montana and south to Califor- nia, Arizona, and New Mexico. This species forms hybrids with Cowania mexicana; such plants are distinguished by the more lobed leaves, and longer and more numerous ach- enes; 122 (xi). Pyracantha Roem. Evergreen shrubs, armed with thorns; leaves alternate, simple, crenate-serrate, pe- tiolate; stipules minute, caducous; flowers perfect, regular, borne in simple or branched corymbs; sepals 5; petals 5, white; stamens 20, the filaments subulate; pistil 1, the ovary inferior, 5-loculed; styles usually 5; fruit a pome. Pyracantha coccinea Roem. Fire-thorn. Shrubs to 3 m tall or more, with thorns 0.5-1.5 cm long or more; leaf blades 0.8-4.5 cm long, 0.5-1.8 cm wide, elliptic to oblan- ceolate, acute apically, cuneate to acute ba- sally, crenate-serrate; petioles puberulent; in- florescence pilosulous at anthesis; sepals broadly triangular; petals white, 3-4 mm long, 2-3 mm wide; pomes red orange, per- sistent. The fire-thorn is a common plant in cultivation through much of Utah; the plants persist following cultivation and escape rarely; Utah Co.; introduced from Eurasia; 6 (0). ' Pyrus L. Trees with unarmed branches; leaves alter- nate, simple, not or seldom lobed; stipules de- ciduous; flowers perfect, regular, borne in co- rymbs; hypanthium short; sepals 5, persistent; petals 5, white; stamens many; pistil 1, the ovary inferior, usually 5-loculed; styles 3-5, separate to the base; fruit usually pear shaped, the flesh with stone-cells. Pyrus communis L. Common pear. Small trees to 6 m tall, the branchlets commonly glabrous; leaves 2-8 cm long, ovate to oblong or elliptic, glabrous or glabrate, leathery, cre- nate-serrulate to subentire; flowering with the leaves; petals white, mainly 12-18 mm long; fruit pear shaped to almost spherical. The pear of commerce is widely grown in Utah, with Bartlett being the most common cultivar; 8 (ii). Rosa L. Shrubs, deciduous; stems armed with prickles or spines, rarely unarmed; leaves al- ternate, pinnately 3- to 9-foliolate; stipules conspicuous, adnate to petioles; flowers per- fect, solitary or in corymbs; hypanthium urn shaped to globose or ellipsoid, red, red or- ange, yellow, or purplish, fleshy at maturity; sepals 5; petals 5 (or numerous in double forms); stamens numerous, inserted on mar gin of ringlike disk; pistils few to numerous; styles exserted through or to orifice of disk; March 1982 Welsh: Utah Flora, Rosaceae 39 fruit of achenes, enclosed in the fleshy hypan- thium (hip). Tlie cultivated roses are largely of hybrid derivation, and are not well repre- sented in herbaria. The key includes both in- digenous and cultivated taxa because of the propensity of some species to escape and of others to persist for long periods following cultivation. The key is tentative at best, be- cause of the propensity of all roses to hybridize. 1. Stipules deeply fringed or pectinate, appearing as lateral projections of petiole base- flowers 'white (or pink in some hybrids keying here); cultivated and ' . „ R. multiflora escapmg ■> - Stipules entire, or rarely fringed, but not cut to the petiole; flowers variously colored 2(1) Flowers mostly 5-9 cm broad; styles long-exserted from hypanthium; cultivated and persisting ^- odorata Flowers mostly less than 5 cm broad; styles not exserted, forming a dense headlike stopper in orifice of hypanthium ^ 3(2). Leaflets stipitate-glandular beneath; sepals strongly stipitate-glandular, erect or spreading in fruit; cultivated and escaping «• ruhiginosa - Leaflets not at all or only sparingly stipitate-glandular beneath; sepals various in fruit; cultivated and escaping or indigenous 4 4(3). Sepals reflexed and finally deciduous after flowering; stipitate glands sparse on lower midveins and sepals; cultivated and escaping ^- camna Sepals erect and persistent following flowering; stipitate glands rarely present; plants indigenous 5(4). Sepals 1.5-4 long; petals 2.2-4 cm long; hips 1-2 cm long at maturity R. nutkana Sepals 1-2.2 cm long; petals 1-2.5 cm long; hips 0.6-1.5 cm long at maturity ... ^ R. woodsii Rosa canina L. Dog Rose. Shrub 1-3.5 (4) m tall; stems sometimes clambering, armed with scattered, strongly curved to straight spines and no prickles, glabrous; stipules en- tire; leaves 3-8 cm long or more, with 3-7 leaflets, the terminal leaflet 0.6-2.2 cm long, 0.3-1.2 cm broad, glabrous to sparingly vil- lous and stipitate-glandular along the veins beneath, serrate to doubly serrate; flowers single, solitary or 2-5 on short glabrous or glandular pedicels; sepals 1.2-1.5 cm long, sparingly stipitate-glandular, reflexed in fruit, soon shattering; petals white or pink. 1-1.8 cm long; hips ovoid, about 1 cm thick, red. Cultivated, persisting, and rarely escaping in lower elevation portions of Utah, specimens examined from Utah and Salt Lake counties; introduced from Europe. Rosa multiflora Thunb. Multiflora Rose. Shrub to 2 or 3 m tall or more; stems some- times clambering, armed with prickles or rarely unarmed, glabrous; stipules about half as long as the petiole, pectinate, cut almost to petiole; leaves 4-9 cm long or more, 5- to 11-foliolate, the terminal leaflet 1.5-3 cm long, glabrous or puberulent and rarely with some glands beneath, serrate; flowers numer- ous (to 50 or more), white (pink in some hybrids), single (double); sepals reflexed in fruit, pubescent; petals 5, or numerous, 0.6-0.8 cm long; hips ovoid, about 6 mm thick, brownish red. Herbarium specimens are very few for this species in Utah, and those involve hybrids with other species and are only tentatively placed herein; Utah Co.; introduced from Japan; 3 (0). Rosa nutkana Presl. Nutka Rose. [R. spal- dingii Crepin]. Shrubs 0.3-2 m tall, rarely more; stems erect or ascending, armed with distinctive infrastipular spines (or rarely unarmed), the internodal prickles lacking or few and different from the infrastipular spines; leaves 6-13 cm long, with 5-7 (9) leaflets, the terminal one 1-7 cm long, 0.8-3.5 cm broad, pubescent to glabrous, rarely stipitate-glandular beneath, serrate to 40 Great Basin Naturalist Vol. 42, No. 1 doubly serrate; flowers solitary (rarely 2 or 3); sepals 1.5-4 cm long, 3-6 mm wide; pet- als 5, pink, 2.2-4 cm long; hips ellipsoid to subglobose, 1-2 cm long and thick, red or- ange to purphsh. Oak-maple, aspen, moun- tain brush, sagebrush, Douglas fir, cotton- wood, and spruce-fir communities at 1525 to 3355 m in Beaver, Box Elder, Carbon, Du- chesne, Garfield, Grand, Juab, Millard, Piute, Salt Lake, Sanpete, Summit, Tooele, and Utah counties; Alaska south to California, Nevada, and Colorado. The features of larger fruit, longer and broader sepals, and usually solitary flowers are diagnostic when taken in combination for most specimens. However, intermediate specimens between Nootka and Woods rose are known. Our material has been assigned to var. hispida Fern.; 48 (vii). Rosa odorata Sweet. Tea Rose; Hybrid Rose. Shrubs, 0.3-6 m long or more; stems erect or ascending to clambering, armed with infrastipular and/or internodal spines or prickles, or unarmed; leaves 4-20 cm long, with usually (3) 5 leaflets, the terminal 1-7 cm long, 0.8-4 cm wide, once or twice ser- rate, glabrous or pilose and somewhat stipi- tate glandular; flowers solitary or usually few to numerous; petals 5, or more commonly nu- merous, of various colors and sizes; hips vari- ous in size, shape, and color. Included within this catchall name are the hybrid tea roses of commerce, grown widely in Utah for orna- ment. The cultivars are mainly complex hybrids, with ancestors involving R. mos- chata J. Herrmann, R. foetida J. Herrmann, R. gallica L., R. chinensis and R. multiflora Thunb. (see Flora Europaea, 1968, p. 26 for a more complete discussion). The hybrid roses persist and are present arotmd abandoned farmsteads in Utah; introduced from Eurasia- 3(i). Rosa rubiginosa L. Sweetbriar. [R. eglan- teria L.]. Shrubs 0.5-2 m tall; stems erect or ascending, armed with distinctive flattened infrastipular spines and often with straight in- ternodal prickles; leaves 3.5-10 cm long, with 5-7 (9) leaflets, the terminal one 1.2-3.5 cm long, 0.8-3 cm wide, conspicuously stipi- tate glandular on one or both surfaces; doubly serrate; flowers solitary or 2-4; sepals 1-2 cm long; petals 0.8-2 cm long, pink to white; hips 1-1.5 cm long, ellipsoid to sub- globose. Cultivated, persisting, and escaping in Salt Lake and Tooele counties, and likely elsewhere in Utah; introduced from Utah- 2 0. Rosa woodsii Lindl. Woods Rose. [R. fend- leri Crepin; R. neomexicana Cockerell; R. manca Greene; R. arizonica Rydb.; R. cali- fornica var. ultramontana Wats.; R. macounii Greene; R. chrysocarpa Rydb., type from Abajo Mts.; R. puberulenta Rydb., type from Montezuma Canyon]. Shrubs 0.1-2.5 (3) m tall; stems armed with infrastipular spines and/or internodal prickles or spines, or unarmed; leaves 1.5-13 cm long, 0.4-2.5 cm broad, pubescent to glabrous and rarely stipi- tate glandular, serrate to doubly serrate; flowers solitary or 2 to several; sepals 1-2.2 cm long, 2-3.5 cm wide; petals 5, pink (rarely white), 1-2.5 cm long; hips ellipsoid to subglobose, 0.6-1.5 cm long and thick, red orange to yellow. Streamsides, irrigation canals, marsh lands, lake shores and hillsides in palustrine, lacustrine, and riparian habi- tats; also in mountain brush, juniper, aspen, and spruce-fir communities at 850 to 3265 m in all Utah counties; Alaska and MacKenzie east to Hudson Bay and south to California, Texas, Missouri, and Wisconsin. The species is represented in Utah by a variable assem- blage that has been included within the con- cept of var. ultramontana (Wats.) Jeps. There are plants from Garfield and San Juan coun- ties especially, which have very coarse inter- nodal as well as infrastipular spines. These would key to R. neomexicana Cockerell, but gradient specimens tie these striking excep- tions to the mass of variation within the com- plex of forms. The striking species R. stellata Woot. might occur in southern Utah. The young stems of that species are stellate- pubescent or copiously glandular hispidulous or both, and the older stems are armed with numerous long, nearly straight prickles; 140 (XV). RuBus L. Shrubs; stems armed with prickles or bristles, or unarmed; leaves alternate, pin- nately compound or palmately veined and lobed; stipules various, usually persistent; flowers perfect or imperfect, regular, solitary or few to numerous in cymes; hypanthium March 1982 Welsh: Utah Flora, Rosaceae 41 short, saucerUke, Uned with a glandular disk; sepals usually 5, lacking bracteoles; petals the same number as the sepals; stamens 15 to nu- merous, linear-subulate; pistils several to many, the ovaries superior, each 1-loculed; styles 1 per pistil, the stigma capitate; fruit of separate drupelets, or the drupelets coherent and free of the receptacle, hence an "aggre- gate" fruit. Bailey, L. H. 1941. Species Batorum. The genus Rubus in North America. Gentes. Herb. 5: 1-932. 1. Leaves simple, palmately veined and lobed, green on both sides; stems unarmed ^ — Leaves compound pinnately 3 to 5 foliolate; stems armed 3 2(1). Leaves mainly less than 6 cm wide, the lobes rounded in general outline, green above, white-tomentose beneath; flowers mainly solitary, plants rare, known only from low elevations in San Juan Co R- neomexicana — Leaves mainly 6-30 cm wide, the lobes acute to attenuate in general outline; flowers borne in clusters of 2-6 or more plants locally common, montane, widespread R. parviflorus 3(1). Main prickles straight, slender, retrorsely disposed along stem; fruit red when j.jpg R. idaeus — Main prickles flattened, curved or straight, retrorse or retrorsely curved 4 4(3). Receptacle fleshy, the drupelets adhering, not slipping free when ripe; stems usually strongly armed, trailing or clambering; cultivated, persisting, and escaping R. discolor — Receptacle not fleshy, the drupelets slipping free when ripe; stems arching, but not trailing or clambering; indigenous and with cultivated phases K. leucodermis Rubus discolor Weihe & Nees Himalayan Blackberry. [R. procerus Muell.-Arg.]. Shrubs, often clambering or sprawling, the stems to several m long, armed with strong, straight, flattened spines; stipules linear, entire; leaves 7-20 cm long, pinnately to palmately com- pound, with 3-5 leaflets, the terminal leaflet 3-12 cm long, 2-8 cm wide, green and glabrous above, tomentose beneath; flowers usually perfect, conspicuous, mainly 3-20 in clusters; sepals 6-10 mm long, lanceolate; petals white (rarely reddish), 10-15 mm long; staminal filaments linear; pits to 3 mm long, the drupelets adherent to receptacle, numer- ous, the flavor agreeable. Roadsides, field margins, and abandoned farmsteads in Juab, Utah, and Washington counties (likely else- where); introduced from the Old World; 3 (i). Rubus idaeus L. Raspberry [R. strigosus Michx.; R. idaeus var. strigosus (Michx.) Max- im.; R. sachalinensis H. Levi.; R. idaeus ssp. sachalinensis (H. Levi.) Focke; R. idaeus var. canadensis Richards.; R. melanolasius Dieck]. Shrubs, 2-15 (20) dm tall, the stems, petioles, and veins on lower leaf surfaces with glandular pricklelike processes or prickles, or both; stipules linear; leaves 2-20 cm long, pinnately compound with 3-5 leaf- lets, the terminal leaflet 1.2-10 cm long, 0.6-7.5 cm broad, green and glabrous to hairy above, white or gray hairy to glabrate and greenish beneath; flowers perfect, not conspicuous, solitary or 1 to few in clusters; sepals 4-12 mm long, lanceolate; petals white, 4-7 mm long; staminal filaments slen- der, often somewhat clavate; pits 2-2.5 mm long, the drupelets coherent, red, several to many, the flavor agreeable. Riparian sites and talus slopes in aspen and mixed conifer com- munities at 2135-3420 m in Beaver, Carbon, Duchesne, Emery, Garfield, Grand, Iron, Juab, Kane, Piute, Salt Lake, Sevier, Summit, Tooele, Uintah, Utah, Wasatch, and Wash- ington counties; Alaska east to the Atlantic and south to California, Mexico, Iowa, and North Carolina; Eurasia. Our indigenous ma terial belongs to ssp. melanolasius (Dieck) Focke. Cultivated phases belong mainly to ssp. idaeus; 48 (xiii). 42 Great Basin Naturalist Vol. 42, No. 1 Rubus leucodermis Dougl. ex T. & G. Black Raspberry. Shnibs, mainly 1-3 m long, the stems, petioles and some veins on lower leaf surfaces armed with retrorsely curved, flattened, catclawlike prickles; stipules lin- ear; leaves 6-14 cm long, pinnately com- pound, with 3-5 leaflets, the terminal leaflet 3-7.5 cm long, 2-6 cm wide, green and al- most or quite glabrous above, white-tomen- tose beneath; flowers usually perfect, not conspicuous, mainly 2-10 in clusters; sepals 6-12 mm long, lance-acuminate; petals white, shorter than the sepals; staminal fila- ments slender, linear-subulate; pits to 2.5 mm long, the drupelets coherent, several to many, the flavor agreeable. Dry open slopes in mountain brush and in riparian communities at 1678 to 2200 m in Millard, Salt Lake, and Utah counties; British Columbia east to Mon- tana and south to California and Nevada. The plants are rare in collections, and the distri- bution is probably wider than indicated; 4 (0). Rubus neomexicanus Gray. Shrubs, 0.5-1.5 m tall, the stems, petioles, and leaves unarmed, merely villous-puberulent and sometimes minutely glandular; stipules lance- ovate, entire or serrate; leaves palmately lobed and veined, simple, the blades 1.2-4.2 cm long (from sinus to apex), 1.5-5.5 cm wide, green above, pale green beneath, pu- berulent on one or both sides; flowers usually perfect, solitary, showy; sepals 10-14 mm long, lance-ovate, entire or serrate; petals white, 2-17 mm long, the drupelets not espe- cially coherent, red, several to many, thinly fleshed, hardly palatable. Hanging garden with Ostrya knoivltonii, at 1130 to 1160 m in Ribbon Canyon, San Juan Co., and to be sought in other shaded moist alcoves along Lake Powell; Arizona and New Mexico. This is a truly attractive species, with startling large white roselike flowers; 5 (iii). Rubus parviflorus Nutt. Thimbleberry. Shrubs, 0.5-2 m tall, rarely taller, the stems, petioles, and leaves unarmed, stipitate- glandular; stipules lanceolate, entire or ser- rate; leaves palmately lobed and veined, simple, the blades 4.5-15 cm long (from sinus to apex), 5.5-20 cm wide, green above, pale beneath, pubenilent on one or both sides, or glabrate above; flowers usually perfect, in clusters of 2-7, showy; sepals 8-19 mm long. ovate, the apex caudate-attenuate, entire; petals white, 13-18 (20) mm long, or more; staminal filaments linear-subulate; pits to 3 mm long, the drupelets coherent as an aggre- gate, red, numerous, thinly fleshy, almost dry at maturity, palatable. Riparian habitats in aspen, spruce, fir, lodgepole, Douglas fir, mountain brush at 1435 to 2745 m in Du- chesne, Salt Lake, San Juan, Sanpete, Sum- mit, Tooele, Utah, Wasatch, and Weber counties; Alaska east to Great Lakes, and south to California, Arizona, New Mexico, and the Dakotas. Our plants belong to var. parviflorus; 25 (i). Sanguisorba L. Perennial herbs, from a branching caudex; leaves basal and alternate, pinnately com- pound; stipules adnate to the petioles, per- sistent; flowers mostly imperfect, regular, nu- merous in short to elongate, dense spikes; hypanthium subglobose, restricted near the apex; sepals 4, petaloid; petals lacking, sta- mens numerous; pistils 1-3, the ovary superi- or, 1-loculed; styles 1 per pistil, the stigma capitate, fringed; fruit an achene, enclosed by the usually 4-angled to 4-winged hypanthium. Sanguisorba minor Scop. Burnet. Plants mainly 2-5 dm tall; caudex clothed with per- sistent stipules and petioles; basal leaves 4-18 cm long, with mostly 9-17 oval to obovate- oblong leaflets, 0.6-1.8 cm long, coarsely ser- rate; spikes subglobose to cylindroid, 8-40 mm long; bractlets ovate; flowers mainly im- perfect, the lower staminate and the upper pistillate; calyx greenish or pinkish; hypan- thium cone shaped in fruit, woody; stamens numerous, the filaments filiform, long-ex- serted. Introduced revegetation and erosion control plant at 1525 to 2135 m elevation in Garfield, Tooele, Utah, and Washington counties; introduced from Europe; 7 (ii). SiBBALDIA L. Perennial herbs from a caudex; leaves basal or cauline and alternate, long-petioled, pal- mately 3-foliolate; stipules adnate to petioles, persistent, lanceolate; flowers perfect, regu- lar, borne in leafy-bracted cymes; hypan- thium short, saucer shaped, lined with a March 1982 Welsh: Utah Flora, Rosaceae 43 glandular disk; sepals 5, alternating with 5 se- paloid bracteoles; petals 5; stamens usually 5; pistils 5-20, distinct, the ovaries superior; styles 1 per pistil, the stigmas capitate; fruit an achene. Sibbaldia procumbens L. Sibbaldia. Plants low, mat forming, the flowering stems 0.4-1.4 dm tall; leaves 2-12 cm long, the 3 leaflets oblanceolate to obovate, 3 (rarely 5)- toothed apically, the terminal leaflet 11-32 mm long, 7-18 mm broad, stiffly hairy on both surfaces; flowers inconspicuous; sepals 2.5-5 mm long; petals pale yellow, 1.5-3 mm long; achenes stipitate, about 1 mm long. Al- pine timdra, krumholz, spruce-fir, meadow, and lodgepole pine communities, often in talus or gravel at 2745 to 3660 m in Beaver, Daggett, Duchesne, Grand, Piute, Summit, Uintah, Utah, and Wayne counties; Alaska east to Newfoundland and south to Califor- nia, Colorado, Quebec, and New Hampshire; circumboreal; 28 (vi). Sorbaria (Ser.) A. Br. Sh-ubs with unarmed branches; leaves al- ternate, pinnately compound; stipules per- sistent; flowers perfect, regular, borne in ter- minal panicles; hypanthium short, lined with a glandular disc; sepals 5, persistent; petals 5, white; stamens 20-50; pistils 5, somewhat connate basally; styles 1 per pistil, the stigmas capitate; fruit of follicles. Sorbaria sorbifolia (L.) A. Br. Sorbaria. Shrubs to 2 m tall, sometimes taller; leaves 8-20 cm long or more; leaflets 11-23, lan- ceolate to oblong-lanceolate, serrate or doubly so, long-acuminate, glabrous or pu- berulent, the hairs stellate; inflorescence 10-25 cm long; flowers white, about 8 mm wide; hypanthium glabrous; fruit glabrous. Cultivated ornamental in Davis, Salt Lake, and Utah counties, and probably elsewhere; introduced from Asia; 4 (0). SORBUS L. Shrubs or small trees with unarmed branches; leaves alternate, pinnately lobed or compovmd; stipules persistent or deciduous; flowers perfect, regular, numerous in corym- bose cymes; hypanthium short, lined with a glandular disk; sepals 5, persistent; petals 5, cream to white; stamens 15-20; pistils 1, the ovary inferior, 2- to 5-loculed; styles 2-5, the stigmas capitate; fruit a pome. Jones, G. N. 1939. A synopsis of the North American species of Sorhus. J. Arnold. Arb. 20: 1-43. 1. Leaves simple, lobed or pinnatifid; petioles and branchlets of inflorescence densely white villous-tomentose S. hybrida - Leaves compound; petioles and branchlets of inflorescence sparingly tomentose 2 2(1). Winter buds densely white- villous, the surface obscured by the hairs; maximum leaflet number commonly 15; plants cultivated S. aucuparia — Winter buds sparingly hairy, the shiny surface not at all obscured by the hairs; maximum leaflet number 13; plants indigenous S. scopulina Sorbus aucuparia L. European Mountain- ash. Trees, mostly 3-6 m tall, with grayish or yellowish green smooth bark; winter buds densely white-villous; leaves pinnately com- pound; leaflets 11-15, 3-5 cm long, 1-1.8 cm broad, the margins coarsely serrate except at the base; petioles and branches of in- florescence sparingly white-hairy at least in flower; stipules persistent; flowers 8-10 mm broad; sepals triangular; petals white to cream, orbicular, 3-4.5 mm long; fruit 9-11 mm long, scarlet, drying purplish. Cultivated ornamental, persisting, and escaping (?) in Salt Lake, Summit, and Utah counties; in- troduced from Europe; 11 (0). Sorbus hybrida L. Trees, mostly 3-6 m tall, with grayish or yellowish green smooth bark; winter buds white-villous-tomentose; leaves simple or pinnatifid, usually with at least one pair of lobes free at base of blade, the lobes coarsely serrate or doubly serrate; petioles and branches of inflorescence dense- ly white, villous-tomentose; stipules de- ciduous; flowers 10-14 mm broad; sepals 44 Great Basin Naturalist Vol. 42, No. 1 triangular; petals white to cream, broadly el- liptical 5-6 mm long; fruit 10-12 mm long, globose, red. Cultivated ornamental, per- sisting, in Utah county; introduced from Eu- rope; 7 (0). Sorbus scopulina Greene. Shrubs, 1-4 m tall, with grayish-red or yellowish bark; win- ter buds glutinous and glossy, white-hairy to glabrous; leaves pinnately compound; leaflets 7-13, 2-9 cm long, 0.7-3 cm broad, sharply serrate almost to the base; branches of in- florescence sparingly to rather densely pu- bescent with white hairs; stipules persistent or tardily deciduous; flowers 8-12 mm broad; sepals triangular; petals white to cream, oval, 4-6 mm long; fruit 5-10 mm long, scarlet to orange, drying purplish. Aspen, spruce-fir, white fir, Douglas fir, and ponderosa pine communities at 2075 to 2900 m in Carbon, Duchesne, Salt Lake, San Juan, Sanpete, Summit, Utah, Wasatch, Washington, and Weber counties; Alaska south to California, New Mexico, and the Dakotas; 28 (vi). Spiraea L. Deciduous shrubs with unarmed branch- lets; leaves alternate, simple; stipules obso- lete; flowers perfect, regular, borne in termi- nal corymbs; hypanthium cup shaped; sepals 5, persistent; petals 5; stamens 25 or more; pistils 3-7 (usually 5), distinct, the ovaries su- perior, each 1-loculed; styles 1 per pistil, the stigmas capitate; fruit a few-seeded follicle. Spiraea x vanhouttei (Briot) Zabel. Shrubs to 2 m tall; stems finally arching; leaves 0.8-3.5 cm long, 0.4-1.7 cm wide, cuneate- obovate, serrate to doubly serrate at the apex and often 3- to 5-lobed; inflorescences pe- dunculate, terminal on short lateral branches; petals white, 3.5-4.5 mm long, oval; follicle to 5 mm long (including styles). Commonly cultivated ornamental, persisting in Carbon, Salt Lake, and Utah counties; introduced from Eurasia. This plant a hybrid involving S. cantoniensis Lour, and S. trilohata L.; 5 (0). SEASONAL FOODS OF COYOTES IN SOUTHEASTERN IDAHO: A MULTIVARIATE ANALYSIS James G. MacCracken^-^ and Richard M, Hansen' Abstract.- Seasonal foods of coyotes (Canis latrans) inhabiting the Idaho National Engineering Laboratory site were examined using step-wise discriminant analysis. Significant differences (P < 0.01) were detected among seasons in food consumption by coyotes, where univariate statistical analysis failed to recognize differences. Recognition of seasonal changes in foods consumed by coyotes is essential to understanding coyote feeding strategies. The role op- portunistic behavior plays in coyote food selection on the study area is questioned. Coyotes {Canis latrans) have been and con- tinue to be a center of controversy (Bailey 1907, Taylor et al. 1979). As a result, numer- ous studies have been published dealing with many aspects of coyote ecology (Bekoff 1978). Food habits of coyotes are well docu- mented in the literature for a variety of eco- logical conditions (Murie 1935, Clark 1972, MacCracken 1981). Few studies, however, have evaluated coyote foods on a seasonal basis (Meinzner et al. 1975). Although many studies have shown that one or two items make up the bulk of coyote foods (Sperry 1941, Murie 1945, Gier 1968, Johnson and Hansen 1979a, and others), the relative abun- dance of these prey species experiences sea- sonal fluctuations. The seasonal availability and abundance of some food items would presumably result in seasonal differences in coyote diets if coyotes are truly opportunistic feeders. Johnson and Hansen (1979a) and MacCracken (1981) both questioned the de- gree opportunistic behavior plays in coyote feeding. In the past, discussion of seasonal differ- ences in coyote foods has largely been based on observed changes of relative amounts of a single item in a coyote dietary. Statistical analysis has been limited due to the number of variables (food items) involved, violation of assumptions of univariate tests, and the lack of a test's power in detecting differences. The purpose of this paper is to present data on seasonal coyote foods in southeastern Idaho and to discuss the application of a mul- tivariate procedure in detecting differences in coyote food selection. Study Area and Methods This study was conducted on the Idaho Na- tional Engineering Laboratory (INEL) site in southeastern Idaho from October 1977 through July 1979. Johnson and Hansen (1979a) studied coyote food habits on the INEL site from July 1975 to July 1977. The INEL site was located on the Upper Snake River Plain, which is dominated by sagebrush-grass vegetation associations (Har- niss and West 1973, Anderson and Holte 1981). Eggler (1941) gave a description of the geology and climate of the plain. The INEL site was divided into two areas based on the presence or absence of coyote control programs. Control activities were confined to the peripheral portion of the INEL site. Livestock grazing was also con- fined to the peripheral portion of the study area. Coyote feces were collected from 25 per- manent transects systematically located as described by Johnson and Hansen (1979a). We included transitional areas defined by Johnson and Hansen as being part of the cen- tral area in this study. Transects were 'Department of Range Science, Colorado State University. Fort Collins. Colorado 80523. 'Present address: USDA, Forest Service, Rocky Mountain Forest and Range Experiment Station, Rapid City, South Dakota 57701. 45 46 Great Basin Naturalist Vol. 42, No. 1 gleaned of all coyote feces in July 1977. Coy- ote feces examined in this study were collect- ed in October 1977 and 1978, representing summer diets, in December 1977, represent- ing fall food consumption, June 1978 and April 1979, representing primarily winter diets, and July 1979, which represented a pe- riod of spring feeding by coyotes. Coyote feces were dried at 60 C for 48 hours in a forced-air-drying oven, then weighed. Each dried scat was placed in a fine mesh nylon bag, soaked for 24 hours in tap water, then cleared of all soluble material by agitating in a clothes washer. After all soluble material had been removed, scats were tum-- bled dry in a clothes drier. Food items in scats were identified by comparison with reference materials and re- corded by frequency of occurrence. Frequen- cy of occurrence of food items was converted to grams of dry matter ingested following procedures explained by Johnson and Hansen (1979b). An estimate of coyote food consumption was determined for each of the 25 transects for each collection date. Five diet estimates were randomly selected for analysis, using a table of random digits (Snedecor and Coch- ran 1967) for each collection date and both areas of the INEL site, which included ap- proximately 550 feces. Differences in coyote food consumption among seasons were tested for significance with step-wise discriminant analysis (Hope 1968, Cooley and Lohnes 1971, Klecka 1975). Discriminant analysis determined which vari- ables (food items) were the most useful in dis- tinguishing between seasons, developed equa- tions (discriminant functions) that classified diet estimates as to season of feeding, and in- dicated which variables contributed the most information to a particular function. Results Nuttall cottontails (Sylvilagus nuttali), montane voles {Microtus montanus), and northern pocket gophers (Thomomys tal- piodes) made up the bulk of coyote foods during the period of this study (Table 1). Sig- nificant differences (P < 0.01) were detected among seasons in coyote food consumption as each variable was entered into discriminant analysis. All seasonal diets were different (P < 0.01) after 15 of 21 food items had been considered. Those 15 food items were the Table 1. Mean (± SE) percent relative dry weight (g) of food items recovered from coyote feces collected on a seasonal basis from the Idaho National Engineering Laboratory. Number of diet estimates is in parentheses.' Season Food items Winter(12) Spring(14) .Summer(22) Fall(12) Mammals Sylvilagus nuttalli 32 ± 8 40 ± 8 69 ± 5 61 ± 6 Brachylagus idahoensis 4 ± 4 13 ± 5 2 ± 1 2 ± 2 Lepiis californicus 9 ± 5 ° 1 ± 1 4 ± 2 L. townsendi 1 ± 1 2 ± I Microtus montanus 12 ± 5 16 ± 6 11 ± 3 7 ± 3 Thornoinys talpoides 3 ± 2 11 ± 4 4 ± 2 4 ± 2 Perognathus sp. 1 ± 1 1 ± 1 5 ± 2 1 ± 1 Cricetid mice 3 ± 2 1 ± 1 1 ± 1 7 ± 4 Antihcapra americana 3 ± 2 1 ± 1 1 ± 1 7 ± 3 Livestock 11 ± 6 1 ± 1 Spennophilus townsendi 1 ± 1 9 ± 3 2 ± 1 Dipodornys ordi 6 ± 4 1 ± 1 » 1 ± 1 Eutamias minimus 2 ± 1 2 ± 2 Mannota flaviventris 3 ± 3 Neotoma cinerea 1 ± 1 Birds 1 ± 0 2 ± 1 1 ± 1 . Reptiles ° 2 ± 1 o Insects 4 ± 4 1 ± 0 1 ± 0 o Seeds 0 o o Plant fragments 1 ± 0 o . 1 ± 0 Grass macrofragnients 5 ± 5 ° ' •XI .X4 . 'xe yV P3 'XZ •X3 ^'^ H4 'Gl •HI G2 H2^ 0 20 40 60 80 100 X-AXIS Fig. 2. Ordination of all 18 sites studied, using all spe- cies sampled. Literature Cited Beals, E. W. 1973. Ordination: mathematical elegance and ecological naivete. Journal of Ecology 61:23-35. Bell, K. L., and L. C. Bliss. 1979. Autecology of Ko- bresia bellardii: why winter snow accumulation limits local distribution. Ecological Monographs 49:377-402. Billings, W. D. 1979. High mountain ecosystems. Evo- lution, structure, operation, and maintenance. Pages 97-125 in P. J. Webber, ed., High altitude geoecology. AAAS Selected Symposium 12. Bliss, L. C. 1963. Alpine plant communities of the Pres- idential Range, New Hampshire. Ecology 44:678-697. Bray, J. R., and J. T. Curtis. 1957. An ordination of the upland forest communities of southern Wiscon- sin. Ecological Monographs 27:325-349. Cronquist, a., a. H. Holmgren, N. H. Holmgren, J. L. Reveal, and P. K. Holmgren. 1977. Inter- mountain flora: vascular plants of the Inter- mountain West. Vol. 6. Columbia Univ. Press, New York. 584 pp. Donahue, R. L., J. C. Shickluna, and L. S. Robertson. 1971. Soils: an introduction to soils and plant growth. Prentice-Hall, Englewood Cliffs, New Jersey. Flock, J. W. 1978. Lichen-bryophyte distribution along a snow-cover-soil-moisture gradient, Niwot Ridge, Colorado. Arctic and .'Alpine Research 10:31-45. Gleason, H. a. 1939. The individualistic concept of the plant association. .'American Midland Naturalist. 21:92-110. Hayward, C. L. 1952. Alpine and biotic communities of the Uinta Mountains. Ecological Monographs 22:93-118. Holmgren, A. H., and J. L. Reveal. 1966. Checklist of the vascular plants of the Intermountain Region. U.S. Forest Service Research Report INT-32. In- termountain Forest and Range Experiment Sta- tion, Ogden, Utah. 160 pp. HoLWAY, J. C, AND R. T. Ward. 1963. Snow and melt- water effects in an area of Colorado alpine. American Midland Naturalist 69:189-197. Johnson, P. L., and W. D. Billings. 1962. The alpine vegetation of the Beartooth Plateau in relation to cryopedogenic processes and patterns. Ecological Monographs 32:105-135. Lewis, M. E. 1970. Alpine rangelands of the Uinta Mountains, Ashley and Wasatch national forests. Table 5. R values for simple linear regressions ran between ordination axes and site parameters. An asterisk (°) indicates significance at the 0.05 level. All 18 sites Factor All species Important species y Phosphonis Nitrogen Cation exchange pH Water (%) Texture Rocks # Rock size x Rock s/x slope Standing crop Veg. cover Soil cover Rock cover Soil covered by lichen (° Rock covered by lichen .37 .29 .26 .01 .20 .43 .25 .37 .12 .12 .04 .36 .34 .27 .53° ..36 .63» .31 .67° .18 .39 .51° .37 .43 .01 .08 .01 .23 45 .28 .32 -.25 04 M .07 -.27 18 -.67 .17 .41 14 -.82° .11 -.71 38 -.59° .20 -.49 16 .56° .18 .59 38 .29 .17 .44 21 -.05 -.26 -.18 38 .19 .22 .15 March 1982 Briggs, MacMahon: Alpine Plant Communities 59 United States Department of Agriculture, Ogden, Utah. 75 pp. Marr, J. W. 1961. Ecosystems of the east slope of the Colorado Front Range. Univ. of Colorado Studies Series in Biology, No. 8. Univ. of Colorado Press, Boulder, Colorado. 134 pp. MuELLER-DoMBOis, D., AND H. Ellenberg. 1974. Aims and methods of vegetation ecology. John Wiley and Sons, New York. 547 pp. NiMLOs, T. J., AND R. C. McCoNNELL. 1965. Alpine soils of Montana. Soil Science 99::310-321. Scott, D., and W. D. Billings. 1964. Effects of envi- ronmental factors on standing crop and produc- tivity of an alpine tundra. Ecological Mon- ographs 34:243-270. Thilenius, J. F. 1975. Plant production of three high- elevation ecosystems. Pages 60-75 in D. H. Knight, coord.. The Medicine Bow ecology proj- ect, final report, February 28, 1975. Univ. of Wyoming, Laramie, Wyoming, for Division of Atmosphere and Water Resource Management, Bureau of Reclamation, U.S. Dept. Interior, Den- ver, Colorado. Webber, P. J., J. C. Emerick, D. C. Ebert May, and V. Komarkova. 1976. The impact of increased snowfall on alpine vegetation. Pages 201-255 in H. W. Steinhoff and J. D. Ives, eds., Ecological impacts of snowpack augmentation in the San Juan Mountains of Colorado. Colorado State Univ. Press, Ft. Collins. Table 5 continued. 17 sites 13 sites All species Important species All species Important species X v X y X y X y -.14 .14 -..39 .27 -.28 .28 -.76 -..37 .24 -.20 .02 .52° .01 -.01 -.23 .11 .13 .37 -.02 .42 -.21 -.17 .01 .40 .59° -..38 .49 .,35 -.18 .13 .12 .16 .02 -.16 -.08 .50 -.08 .08 -.60° -.23 .62° -..39 .14 ..53° -.10 -.15 .18 .26 .21 .01 .06 -.15 .25 -.15 -.45 .26 -.46 -.07 -.23 -.29 -.17 -.26 ..30 .37 .25 .17 -.,50 -.01 .19 -.29 -.27 -.20 -..39 ..35 .01 .52° .01 .52° .13 .,55° -.23 ..38 -.23 -.85° -.48 -.01 .21 .22 -.36 ..36 -.14 -.72° .09 .22 .32 ..36 -.18 .08 -.05 -.08 -.05 -.08 .58° .37 .26 .08 .26 -.21 .26 -.21 .16 .05 ..35 -.21 -..35 -.17 .01 -.55° -.23 .22 -.24 -.49 .07 -.86° -.23 .28 -.19 .58° OBSERVATIONS ON THE REPRODUCTION AND EMBRYOLOGY OF THE LAHONTAN TUI CHUB, GILA BICOLOR, IN WALKER LAKE, NEVADA James J. Cooper' Abstract.- Various aspects of the reproduction and embryology of Walker Lake Lahontan tui chub, Gila bicobr, were investigated during the spring-summer period of 1976, 1977, and 1981. Tui chub were found to spawn in littor- al regions of the lake beginning in late May or early June. Early in the season male chub substantially outnumbered females over the spawning grounds, with a normal 1:1 sex ratio gradually approached as the season progressed. The developmental period between fertilization to hatch-out was shortened by increases in water temperature. Selected stages of embryonic development are described from egg fertilization through post-hatch. The Lahontan tui chub, Gila bicolor, is a cyprinid found in the Columbia, Klamath, and Sacramento river systems as well as in a nimiber of isolated interior basins of Califor- nia, Oregon, and Nevada (Bailey and Uyeno 1964, Moyle 1976). The Walker, Carson, Truckee, and Humboldt river system of the Lahontan Basin all support populations of tui chub (La Rivers 1962), as well as numerous isolated springs in the north central Great Basin (Hubbs et al. 1974). Little information exists concerning the re- productive behavior and embryology of the tui chub, especially in the Lahontan system, where they are an integral trophic link in the food chain and heavily preyed upon by vari- ous piscivorous fishes. Kucera (1978) reported on the reproductive biology of the tui chub from Pyramid Lake, Nevada. Kimsey (1954) described some early life history information of the tui chub from Eagle Lake, California. Harry (1951), working on the same project produced one of the first papers on tui chub embryology. Spawning and general repro- ductive behavior of tui chub in East and Paulina lakes, Oregon, was described by Bird (1975). Other less quantitative literature on the subject includes observations on Pyramid Lake, Nevada (Snyder 1918, La Rivers 1962). Study Area Walker Lake, the second largest natural body of water entirely within the state is located in west central Nevada approx- imately 209 km southeast of Reno and 10 km north of Hawthorne. The lake is a remnant of pluvial Lake Lahontan, which once occupied west central and northwestern Nevada as well as the Honey Lake region of north- eastern California (La Rivers 1962). Walker Lake is a terminal, alkaline-saline lake with a total dissolved solids content of over 12,500 mg/1, of which sodium chloride, sulfates, and bicarbonates make up approximately 97 per- cent of the total ionic content (Koch et al. 1979). Walker Lake has a surface area of 15,000 ha and a maximum length of 25 km and max- imum width of 9 km. The maximum depth of the lake is near 33 m and the mean depth is 20 m. The elevation of Walker Lake has been declining very rapidly in recent geological time, with subsequent increases in salinity and alkalinity. Since 1915 the level of the lake has dropped at an average rate of 0.58 m per year (Koch et al. 1979). Upstream agri- cultural diversion of the Walker River, the only tributary entering the lake, has been the primary factor responsible for the increased rate of desiccation since the turn of the century. Methods The data presented in this paper were col- lected during the spring-summer period of 1976, 1977, and 1981. Observations of tui 'Bioresources Center, Desert Research Institute, Reno, Nevada 89506. 60 March 1982 Cooper: Lahontan Tui Chub 61 chub spawning in Walker Lake were accom- pUshed by slowly and quietly approaching shore areas during the spawning season. Spawning chub were easily detected as their fins would break the water surface, their presence many times accompanied by splash- ing. Sampling techniques included the use of a 15.2 m beach seine, multipaneled gill nets, and hook and line. Notes on behavior and en- vironmental variables such as water temper- ature, depth of spawning, and substrate type were all recorded concurrently. The embryological development of the tui chub was observed and monitored through an incubation system that closely simulated lake conditions. Two 6.5 1 May-Sloan hatching jars were set up at the Desert Research In- stitute's mobile laboratory on the shore of Walker Lake. Lake water was pumped to a large holding tank on a hill behind the facil- ity and a line was installed to each jar. Each of the lines was equipped with a valve to control flow volumes through the hatching jars at 0.5-1 1 per minute. During the in- cubation periods temperature was monitored at least every four hours with a thermometer measuring to the nearest 0.5 C. Spawn was obtained by capturing ripe males and females in gill nets set near shore. At least two females and five males were used to produce the fertilized eggs for each incubation period. Ripe females were spawned by hand into a flat-bottomed plastic bowl immediately after capture. Milt from ripe males was extruded over the eggs and lake water added. After the water was added, the bowl was swirled gently to mix the sperm and eggs. After the initial mixing, the ferti- lized eggs were allowed to stand for a few minutes and then were thoroughly rinsed with lake water a number of times. Approx- imately one-half hour after water hardening, the eggs were introduced to the hatching jars. Developmental sequences were obtained with the use of a compound Tiyoda micro- scope and a 35 mm Minolta camera with a Vivitar microscope adaptor, using Kodak- Kodacolor ASA 125 film. It was found that micrograph magnification revealed the most detail at 40X. Line drawings were produced from these photographs. Results and Discussion Spawning Tui chub from Walker Lake became sex- ually mature during the spring of their third year, with the exception of a few males that were observed to be mature at the end of the second year. This age is consistent with what Kimsey (1954) found in Eagle Lake, Califor- nia, and Kucera (1978) in Pyramid Lake, Nevada. Male and female tui chub are easily identi- fied to their sex during the spawning season. The most obvious change occurs in the male who become covered with small nuptial tu- bercles. The females undergo a slight en- largement of the anal region and exhibit a marked protrusion of the genital papilla. In both sexes the fins take on a slight reddish coloration. Spawning activity was first observed on 8 June and 20 May in 1976 and 1977, respec- tively, at Walker Lake. The surface water temperature on 8 June was 16.5 C and on 20 May was 13.5 C. Kimsey (1954) found spawn- ing to first occur in Eagle Lake near a tem- perature of 15.5 C. Spawning observations revealed large schools of chub within 1-2 m of the shoreline at a depth of from .25 to 1 m. The dorsal and caudal fins of the fish broke the surface of the water in many instances. Substrate type var- ied from small pebbles to large rocks with small amounts of algae attached to their sur- face in many cases. Actual spawning was not observed, but an examination of a spawning site revealed a number of chub eggs between the rocks and attached to algae. Available literature is consistent in stating that tui chub are inshore spawners. In Eagle Lake, California, mature tui chub were found to migrate from the deeper southern end of the lake to the shallower northern end during the spawning season (Kimsey 1954). La Riv- ers (1962) also made the observation that tui chub congregate in shallow shoreline areas to spawn. Gill net catches at Pyramid Lake re- vealed that 97 percent of the adult benthic tui chub population was inshore in July (Vigg 1978). Sex ratios were calculated from a sample of 852 fish collected over a 15 month period. 62 Great Basin Naturalist Vol. 42, No. 1 The ratio of males to females was 1:1.16, which does not deviate substantially from an assumed 1:1 ratio. Pyramid Lake female tui chub were found to survive to an older age than males (Kucera 1978), which may explain the sex ratio favoring females in Walker Lake. Nevertheless, this ratio deviates dramati- cally during the spawning season. On 29 May 1981, 103 tui chub were captured by seining schools over their spawning ground. Males outnumbered females 84 to 19, which is a sex ratio or 4.4:1. From this group, age III fish were represented by 81.5 percent of the indi- viduals, with the remainder ages II, IV, and V. Gill netting from June through July 1976 at inshore locations also revealed males en- tering spawning grounds earlier than females (Fig. 1). On 15 June males comprised 85.1 percent of the population but decreased to 54.3 percent by 19 July. Kucera (1978) found similar results in Pyramid Lake, where from May to June males increased from 49 to 62 percent. Also of significance are the apparent sexual differences in readiness to spawn at various times in the season. Early in the spawning season (late May) all the males were "running ripe" with sperm, whereas none of the fe- males would extrude their eggs following gentle pressure to the abdominal region. The peak in spawning activity was estimated to occur in mid-July, when the sex ratio again approached normality and a majority of the females in the catches were in a ripe condi- tion. Ripe female chub were difficult to find in gill net sets in late July and early August, although all the males would continue to dis- charge sperm with gentle handling. From a 100-1 90- 80- •- 70H < 60- < .? 50-1 a z < 4QH 30- MALES 85.1% 20- 10- 63.0% 14.9% FEMALES N = 442 5 9.5 3 7.0% 6-15-76 6-24-76 7-6-76 7-19-76 DATE Fig. 1. Percentage of male to female Lahontan tui chub caught in bottom .set gill nets in from 5 to 7 m of water between 15 June and 19 July 1976 in Walker Lake, Nevada. March 1982 Cooper: Lahontan Tui Chub 63 total of 174 females collected from an in- shore gill net set, only 3 individuals (1.7 per- cent) were in a running ripe condition on 4 August. Upon examination of apparently spent females, it was found that they still contained eggs, a fact that suggests multiple and protracted spawning. It appears as though males are the first to become sexually active in the spring and the last to become inactive at the termination of the spawning season. Reproductive behavior such as this would seem to ensure many spawning males for each female and in turn a greater chance of egg fertilization. Embryonic Development Observations on the embryological devel- opment and early larval stages of the tui chub were conducted from mid-July to early August 1976 on Walker Lake. The rate of de- velopment was found to be dependent upon water temperatures (Fig. 2). An increase in temperature from 18.8 to 24.4 C accelerated embryo development by approximately 85 percent. Harry (1951) incubated tui chub eggs from Eagle Lake, California, at 7.2 C for the first 100 hours and then allowed temperatures to vary from 1.1 to 28.9 C. At this variable tem- perature regime the eggs hatched out on the 12th day of incubation. Kimsey (1954) also incubated Eagle Lake tui chub eggs in a quart jar, where the air temperature fluc- ^ 140- CO IT O 130 I Q '20 O QC LU 1 10- Q. rf 100 LU 90 TEMPERATURE (C) Fig. 2. Developmental period for Lahontan tui chub eggs incubated at three mean water temperatures, July and August 1976. Walker Lake, Nevada. Bar equals two standard deviations. tuated from 4.4 to 32.2 C, and found the fry actively feeding in 9 days. In East Lake, Ore- gon, Bird (1975) hatched out tui chub eggs in 192 and 142 hours at 14.6 and 21.9 C, respec- tively. He concluded that highly fluctuating incubation temperatures retarded the devel- opmental period in comparison to stable tem- perature regimes. The ability of tui chub eggs to tolerate wide variations in water temperatures is a valuable survival trait for the species. The eggs incubate in water that is shallow enough to warm up during periods of high solar radi- ation and cool down at night. The short dura- tion of the egg stage is also a highly benefi- cial adaptation because environmental conditions have a shorter period of time to cause mortality at this critical stage of development. Lahontan tui chub eggs flow freely from females when they are in a condition to ac- cept fertilization. It was found that when eggs were forcibly extruded fertilization would not occur. Freshly stripped eggs are approximately 1-1.5 mm in diameter, yel- lowish, opaque, and very adhesive. They are demersal and their specific gravity is consid- erably more than that of Walker Lake water because the eggs quickly sink to the bottom. After successful fertilization, the perivitelline space becomes separated from the zygote and swelling occurs, giving the egg as a whole an outer diameter of 1.5-1.8 mm. Stages of embryonic development were monitored continuously from fertilization through hatch-out (Fig. 3). At 21.1 C, by 6 hours the blastodermal cap had formed at the animal pole and covered about one-fourth of the circumference of the yolk. Mid- gastrulation had occurred by 10 hours and late gastrulation by 12.5 hours, with the em- bryo becoming recognizable around the yolk. After 22.5 hours of incubation, the embryo length was approximately two-thirds of the way around the yolk circumference, with the head region discernible from the tail. At 31 hours the embryo had 6-8 somites, the optic vesicles were easily discernible, and the yolk sac was still very large. The embryo began to squirm within the egg case at 44.5 hours and the notochord was visible. At this stage the heart, as well as blood flowing throughout the circulatory system, could be seen. Just 64 Great Basin Naturalist Vol. 42, No. 1 prior to hatching at 96 hours, the embryo had surrounded the egg yolk and completely filled the chorion. At this stage the eyes had become pigmented, and the embryo moved almost constantly. Just after hatching, the lar- va body is curved with slight pigmentation, the swim bladder is visible, and they tend to swim periodically. Afterwards they quickly sink to the bottom. This is consistent with what Bird (1975) found in East and Paulina lakes, Oregon, although Harry (1951) found larvae exceptionally active and able to swim rapidly just following hatching. By 166 hours after fertilization (70 hours after hatching), the larvae were 8-10 mm" long and the pectoral fins had begun to form. Four gill chambers and arches were easily discernible posterior to the eyes, and melano- phores covered more than half of the body. Swimming action had increased but had not yet become consistent. The yolk sac was very small at this stage, but feeding had not yet been observed. The majority of the larvae had died by 237.5 hours, probably of starvation. Most of the fins were present and the body and head were heavily covered with melanophores. Myomeres were well developed and the larva was able to hold itself in a swimming, mid- water position. It is presumed that the fish must soon begin to feed at this stage of devel- opment because the yolk sac had been com- pletely absorbed. Literature Cited Bailey, R. M., and T. Uyeno. 1964. Nomenclature of the tui chub, cyprinid fishes from western United States. Copeia 1964(1): 238-239. Bird, F. H. 1975. Biology of the blue and tui chubs in East and Paulina lakes, Oregon. Unpublished thesis. Oregon State Univ., Corvallis. 165 pp. Harry, R. R. 1951. The embryonic and early larval stages of the tui chub, Siphateles bicolor (Girard), from Eagle Lake, California. California Fish and Game37(2):129-132. HuBBS, C. L., R. R. Miller, and L. C. Hubbs. 1974. Hy- drographic history and relict fishes of the north central Creat Basin. Mem. California Acad. Sci. 7:259. KiMSEY, J. B. 1954. The life history of the tui chub, Siph- ateles bicolor (Girard), from Eagle Lake, Califor- nia. California Fish and Game 40(4):395-410. 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 Cent. Pub. No. 50010. 191 pp. KucERA, p. A. 1978. Reproductive biology of the tui chub, Gila bicolor in Pyramid Lake, Nevada. Great Basin Nat. 38(2):203-207. La Rivers, I. 1962. Fishes and fisheries of Nevada. Ne- vada Fish and Game Comm. 782 pp. MoYLE, P. B. 1976. Inland fishes of California. Univ. of California Press, Berkeley. 405 pp. Snyder, J. O. 1918. The fishes of the Lahontan system of Nevada and northeastern California. Bull. U.S. Bur. Fish. 35:31-86. ViGG, S. 1978. Fish ecology. Chapter 8 in W. F. Sigler and J. L. Kennedy, eds. Pyramid Lake Nevada, ecological study final report. W. F. Sigler and As- sociates, Inc. Reno, Nevada. Fig. 3. Developmental sequences of Lahontan tui chub incubated at 21.1 C. A, One hour after fertilization; B, 6 hours; C, 12.5 hours; D, 22.5 hours; E, 31 hours; F, 44.5 hours; G, 96 hours, just after hatching; H, larva 70 hours after hatching. THE PREVALENCE OF ECHINOCOCCUS GRANULOSUS AND OTHER TAENIID CESTODES IN SHEEP DOGS OF CENTRAL UTAH' Lauritz A. Jensen', Ferron L. Andersen-, and Peter M. Schantz' Abstract.- Fifty-one of 62 sheep dogs in central Utah were successfully purged for diagnosis of cestodes in 1981. Tapeworms were identified in the purged fecal samples of 33 (64.7 percent) animals. Minimum infection rates in the dogs which were purged were; 9.8 percent for Echinococcus granulosus, 29.4 percent for Taenia pisiformis, 27.5 per- cent for T. ovis krabbei, 27.5 percent for T. hydatigena, and 2.0 percent for T. serialis. The prevalence of E. gran- ulosus decreased from 27 percent in 1971 to 9.8 percent in 1981. Echinococcus granulosus is endemic in dogs in central Utah and is primarily con- fined to sheep-raising communities (Andersen et al. 1973). The proportion of infected dogs among those brought to voluntary diagnostic clinics has gradually decreased from 27 per- cent in 1971 (Loveless et al. 1978) to 18 per- cent in 1978 (Condie et al. 1981). In the past, surveys have included all classifications of domestic dogs (e.g., family pet, guard dog, sheep dog, hunting dog, etc.), regardless of the feeding habits. To help determine the ef- fectiveness of the hydatid disease control pro- gram in Utah, and to ascertain if the preva- lence of E. granulosus is in a continued decline, 62 sheep dogs were tested in Sanpete and Summit counties during August and Oc- tober 1981. Field clinics were conducted in the general vicinity of summer range allotments so as to be convenient for sheep herders. Owners were requested to fast their dogs 12 hours prior to the examination. A solution of 1.5 percent arecoline hydrobromide was adminis- tered orally (3 mg/kg of body weight) to in- duce purging, after which the mucoid por- tion of the purge was diluted in water and examined for tapeworms. Specimens of £. granulosus were washed in tap water for 30 minutes and fixed in AFA, whereas the larger taeniids were relaxed in water for 6 hours at ambient temperature and fixed in formalin. Table 1 details the results of the survey. Fifty-one of 62 dogs, which ranged in age from six months to nine years, were success- fully purged. Tapeworms were recovered from the purged fecal specimens of 33 dogs. Infections with E. granulosus were identified in 5 of 51 (9.8 percent) dogs, representing four separate sheep herds. Two of the four owners of these herds previously had had hydatid cysts removed from their liver or lung, and had participated in past field clin- ics. The rates of infection in the 51 dogs were: 29.4 percent for Taenia pisiformis, 27.5 percent for T. ovis krabbei, 27.5 percent for T. hydatigena, and 2.0 percent for T. serialis. The total burden of Taenia in infected dogs ranged from one to 233, with mixed in- fections of two or more species of worms being common. There was no obvious rela- tionship between the age of the parasitized dogs and the proportion of dogs infected, and it was not uncommon for pups, approx- Table 1. Cestodes recovered from 51 sheep dogs of central Utah. No. dogs infected Cestode Sanpete Co. (n = 38) Summit Co. (n=13) Total % dogs infected Echinococcus granidosus Taenia hydatigena Taenia ovis krabbei Taenia pisiformis Taenia serialis 5 5 12 12 1 0 9 2 3 0 9.8 27.5 27.5 29.4 2.0 'Supported in part by NIH Grant Al-10588-10 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Parasitic Diseases and Veterinary Public Health Division, Center for Disease Control, Atlanta, Georgia 30333. 65 66 Great Basin Naturalist Vol. 42, No. 1 imately six months old, to harbor gravid tae- niids. Dogs infected with E. granulosus were injected with praziquantel (Droncit, Bayvet Division, Cutter Laboratories, Shawnee, Kan- sas) at a dosage level of 5 mg/kg of body weight. Comparison of the 9.8 percent rate of in- fection of E. granulosus in 1981 to 27 per- cent in 1971 (Loveless et al. 1978) and 18 percent in 1978 (Condie et al. 1981) suggests a true reduction. This suggestion is further supported by the fact that only dogs at high- est risk, i.e., sheep dogs, were examined in the study herein reported. It is also apparent from the taeniids recovered that sheep herd- ers still feed their dogs ample supplies of sheep viscera, deer, and rabbits. Even though most owners of large sheep herds appear to cooperate and do not give their dogs sheep viscera or wild animals, the transient, hired herders may be less disciplined. Thus, yearly field clinics and educational programs on dis- eases caused by cestodes should be continued. Representative specimens: E. granulosus USNM Helm. Coll. No. 76786; T. pisifomiis No. 76787; T. ovis krabbei No. 76788; T. hy- datigena No. 76789; T. serialis No. 76790. 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. Ainer. Vet. Med. Assoc. 163:1168-1171. 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. Loveless, R. M., F. L. Andersen, M. J. Ramsay, and R. K. Hedelius. 1978. Echinococcus granulosus in dogs and sheep in central Utah, 1971-1976. Amer. J. Vet. Res. .39:499-502. GROWTH OF JUVENILE AMERICAN LOBSTERS IN SEMIOPEN AND CLOSED CULTURE SYSTEMS USING FORMULATED DIETS S. R. Wadley', R. A. Heckmann', R. C. Infanger', and R. W. Mickelsen' Abstract - Growth of juvenile American lobsters, Homarus americanus, raised in four semiopen culture systems, with dailv water exchange rates ranging from 29 percent to 3.3 percent, was compared with growth in a completely closed system. Animals were fed a formulated pelleted ration, water quality factors were mea.sured daily, and changes 'in concentration of nitrate, orthophosphate, and total organic carbon were monitored. Results of two 90-day trials indicate that growth increased in the system with the lower water exchange rates. Maximum growth occurred in the closed system. Interest in commercial culture of the American lobster, Homarus americanus, has increased because of high consumer demand and declining natural fisheries. Dow (1980) indicates that this decline is probably due to overexploitation of the lobster. He states that during the period from 1950 to 1976, inshore landings of lobsters from Newfoundland to New York decreased from 33,000 to 30,000 metric tons and the number of traps used in- creased from 240,000 to 520,000. Other data indicate that in some areas annual trap suc- cess has also declined from 225 pounds per trap in 1889 to 17 pounds per trap in 1970 (John T. Hughes, pers. comm.). The American lobster is one of four marine species given "high priority" status for aqua- culture by the U.S. National Oceanographic and Atmospheric Administration because it meets several requirements for commercial production of aquatic species (Glude 1977). These criteria include adequate consumer de- mand, high profit potential, ability to com- plete the life cycle in captivity, high food conversion efficiency, and resistance to dis- ease (Cobb 1976). The economic potential of lobster culture depends on satisfaction of these factors, plus the development of ef- ficient larval rearing and grow-out systems, production of inexpensive diets, and determi- nation and maintenance of optimum culture environments. The last two factors (formu- lated diets and determination of optimum conditions) are studied in this experiment. Numerous studies have been done on basic nutritional requirements of crustaceans (Gal- lagher et al. 1979, Winget et al. 1976, Castell and Covey 1976). Most researchers agree that formulated diets must replace expensive nat- ural rations for lobster culture to be econom- ically feasible (Van Olst et al. 1980). Conklin et al. (1975) define nutritional requirements for lobsters and report initial development of pelleted rations. The formulation of diets in pellet form is desirable in aquaculture be- cause of ease in production, handling, and feeding (Conklin 1980). Goldblatt et al. (1978) and Infanger et al. (1980) indicate that pelleted rations may lose nutritional quality through vitamin leaching when exposed to culture water. This makes most diets devel- oped thus far unacceptable for lobster culture. Culture systems are usually classified as open, semiopen, or closed. Wheaton (1977) described open systems as production in a natural body of water with few modifica- tions, semiopen systems as those where water is taken from a natural source, passed through the system once and discarded, and closed systems as those where water is placed within the system and is rarely if ever re- placed. The economic production of lobsters is restricted to semiopen and closed systems because natural water temperatures in most areas are below optimal growth temperatures and must be heated to accelerate animal growth. 'Department of Zoology, Brigham Young University. Provo, Utah 84602, 67 68 Great Basin Naturalist Vol. 42, No. 1 Many factors have been used in comparing semiopen and closed systems for culture of marine organisms (King 1973). One impor- tant contrast for lobster culture may be dif- ferences in growth rates between systems. It is unknown if juvenile lobsters will grow faster in a particular type of system. The pur- pose of this experiment was to compare lob- ster growth in five different culture environ- ments ranging from semiopen systems (with four different water exchange rates) to closed systems. Animals in all systems were fed the same formulated diet. Materials and Methods Four semiopen systems and one closed sys- tem were estabHshed in separate 300:1 tanks. Water was removed from each semiopen sys- tem and replaced at different intervals, there- by producing daily water exchange rates of 29 percent in system I, 13 percent in system II, 7.4 percent in system III, and 3.3 percent in system IV. Water in system V remained unchanged during the experiment except for addition of fresh water to compensate for evaporation. A 10 cm thick undergravel filter in each tank provided filtration and a sub- strate for nitrifying bacteria. Culture water was synthetic seawater sim- ilar to that described by Spotte (1970) for large marine aquaria. It was mixed in 1200 liter quantities for use in the experiment. Forty-eight juvenile lobsters were tested in each culture system. Cages in each tank iso- lated animals, which allowed monitoring of the growth rate of each animal throughout the experiment. Two 90-day trials were conducted (trial 1 and trial 2). Growth was determined by mea- suring animal length from eye socket to pos- terior margin of the carapace along the dor- sal midline. Growth of animals between trials could not be compared because lobsters for each trial were hatched from two different females. Comparisons were made between systems within each trial. Test animals were fed a pelleted diet sim- ilar to those described by Infanger et al. (1980). The pellets were produced using a Wenger extruder (model X-5), then sealed in plastic bags and stored at -18 C. Lobsters were fed daily in excess of what could be consumed during the 24-hour period. Food remaining from the previous day was re- moved before feeding a fresh pellet. Water quality was monitored daily in each trial by testing salinity, pH, temperature, dis- solved oxygen, and nitrite. All factors, with the exception of nitrite, were measured di- rectly using the appropriate meter. Nitrite was recorded as percent transmittance, using a Bausch and Lomb Spectronic 20 (Spotte 1970). Results of water quality measurements were compared with optimum levels de- scribed for lobster culture (Van Olst et al. 1980). These optimum levels included a tem- perature range between 20.0 and 22.0 C, 30.0 ppt salinity, 6.4 mg/1 dissolved oxygen, a pH of 8.0, and nitrite levels less than 10.0 mg/1. Spotte (1973) recommends nitrite levels less than 0.1 mg/1 for large marine aquariums. Because nitrite is a potentially toxic waste compound, the lower concentration of 0.1 mg/1 or less was considered optimum. It was determined through use of a standard nitrite solution that a transmittance of 74 percent or higher indicated concentrations lower than this level. Additional tests were performed on water taken from the first" trial by the certified Brigham Young University Environmental Analysis Laboratory. Samples were tested for two elements (copper and iron) and for three compounds (nitrate, orthophosphate, and to- tal organic carbon). These substances may be important toxins or nutrients in culture sys- tems that may strongly affect animal growth (Spotte 1979, Wheaton 1977). Two samples were tested: one sample was freshly made seawater, and the second was the same water taken from system V after the 90-day culture period. Three compounds tested in the first trial (nitrate, orthophosphate, and total organic carbon) were tested more extensively in the second trial from all five water systems. Sam- ples were taken at the beginning of the cul- ture period and again prior to a water change in that particular system. For ex- ample, an initial sample was taken on day 1 from each system and again on day 4 from system I, on day 7 from system II, on day 14 from system III, and on day 30 from system IV. This testing was duplicated twice from March 1982 Wadley et al.: Lobster Growth 69 each semiopen system (series A and B). Wa- ter samples from system V were taken for analysis on days 1, 30, 60, and 90. FIesults Results of juvenile lobster growth experi- ments from both trials include: (1) water quality, (2) detailed water analysis, and (3) growth and survival. Water Quality Water quality was monitored by daily measurement of temperature, salinity, pH, and nitrite. Temperature averaged 20.0 and 21.0 C, mean salinity was 31.0 and 32.0 ppt, pH averaged 8.1, and mean nitrite trans- mittance was 91.5 and 94.3 percent in trial 1 and trial 2 respectively. Dissolved oxygen in trial 1 had a mean concentration of 5.5 mg/1. This value re- mained constant throughout the trial and represents the saturation level at ambient salinity, temperature, and atmospheric pres- sure. Dissolved oxygen was not tested in trial 2. Detailed Water Analysis In comparing 90-day-old water with fresh- ly made synthetic water in trial 1, levels of iron decreased, whereas concentrations of copper, nitrate, orthophosphate, and total or- ganic carbon increased (Table 1). The last three compoimds were tested more exten- sively in trial 2 and compared with concen- trations in all five culture systems. Table 1. Water comparisons: Trial 1. Nitrate Nitrate concentrations increased with the age of culture water within each of the five systems (Fig. 1). Initial levels were below 1.0 mg/1 with the exception of a sample from system I that started at 1.3 mg/1. Final levels ranged from 0.4 mg/1 in system I after 4 days to 11.2 mg/1 in system V after 90 days. The second set of duplicate tests (series B) showed greater increases of nitrate than the first set of tests (series A) in each semiopen system. This probably occurred because samples for the second series were taken later in the cul- ture period, when biological filters were more efficient in converting ammonia to nitrate. Nitrate concentration increased inversely with the amount of water replacement in the four semiopen systems. Final concentrations from the second duplicate series were 1.4 mg/1 in system I after 4 days, 1.9 mg/1 in sys- tem II after 7 days, 5.5 mg/1 in system III af- ter 14 days, and 10.5 mg/1 in system IV after 30 days. Results from system V showed nitrate in- creases over the 90-day culture period, when concentration increased from 0.46 mg/1 on day 1 to 11.2 mg/1 by day 90. Orthophosphate Orthophosphate increased with the age of water within each system (Fig. 2). Initial lev- els ranged from 0.23 mg/1 to 0.41 mg/1, whereas final concentrations ranged between 0.67 mg/1 in system I to 3.40 mg/1 in system Tests New* Ol* Normal levels Copper as Cu (u^/l) 36 49 60C Iron as Fe (ug/1) 92 66 300P Nitrate as N (mg/1) 0.46 2.15 20P Phos-Ortho asP (mg/1) 0.23 2.07 1.0(ug/l)d Total organic carbon (mg/1) 16.66 19.60 6.0d ^Freshly made synthetic seawater '^Synthetic seawater after 90 days '^From Van Gist et al. (1980) '^From Spotte (1979) for marine aquaria Qjy 04 04,07 070 14 0 14 0 300300 306C90 SERIES A BjA ^!' °,* ^ SYSTEM I I M I Ml I IV I V Fig. 1. Nitrate concentration comparisons for systems from trial 2. 70 Great Basin Naturalist Vol. 42, No. 1 V. These levels are greater than concentra- tions found in natural seawater, but compare favorably with concentrations reported in large marine aquaria (Spotte 1979). Total Organic Carbon Levels of total organic carbon (TOC) in- creased with the age of water within each system (Fig. 3). This increase can be seen in system V, which started at 16.7 mg/1 on day 1 and increased to 21.9 mg/1 by day 30, to 27.8 mg/1 by day 60, and to 28.7 mg/1 by day 90. TOC concentration increases did not cor- relate with water exchange when systems - were compared with each other. The greatest differences occurred in system III, which in- creased from 14.7 mg/1 to 37.5 mg/1 in 14 days. The highest final concentration (24.1 mg/1) occurred in system I after only four days. Growth and Survival Results indicate that culture systems do af- fect juvenile lobster growth. In both trials, animals in system V had faster growth rates than any other system (Fig. 4). They were significantly larger (p = .05) than in systems I, II, and III in trial 1 and systems I and II in trial 2 (standard Students t-test). Growth in both trials, with the exception of system II in trial 2, have an inverse relationship with daily water exchange rates even though all differences were not significant. Survival ranged from 46.0 percent to 73.0 percent, averaging 57.2 percent in trial 1, M D*r SERIES SYSTEM and from 50.0 percent to 90.0 percent, aver- aging 72.2 percent in trial 2 (Fig. 5). No clear patterns resulted from either trial. System III had the best survival in trial 2 and the most deaths in trial 1. Discussion Results indicate that water exchange af- fected juvenile lobster growth rates when wa- ter quality factors (temperature, salinity, pH, dissolved oxygen, and nitrate) were main- tained within limits suggested by Van Olst et al. (1980) for lobster culture and by Spotte (1973) for marine aquaria. Growth rates in- creased in both trials with decreasing water exchange, with the best growth occurring in the closed systems. Reasons for this are un- clear, but differences may be partially ex- plained by changes in water chemistry. Wheaton (1977) indicates that water chem- istry will change in culture environments de- pending on the length of time water remains in the system. Sources of the three com- pounds tested in trial 2 (nitrate, orthophos- phate, and total organic carbon) include leached nutrients from unstable diets and waste products from animal metabolism. Copper and iron were not tested in trial 2 be- cause concentrations did not change greatly in trial 1. Increase of nitrate in closed systems is well documented (Liao and Mayo 1972, King SERIES SYSTEM Fig. 2. Orthophosphate concentration comparisons ^'§- ■^- Total organic carbon concentration com- for systems from trial 2. parisons for systems from trial 2. March 1982 Wadley et al.: Lobster Growth 71 1973). Toxicity levels of nitrate are unknown for most species, but Van Olst et al. (1980) suggests 500 mg/1 as the maximum concen- tration desirable for lobster culture. Data from this experiment show increases of ni- trate relative to the age of the water. Levels of nitrate for all systems (highest concentra- tion was 11.2 mg/1 in system V) were well below 500 mg/1, which probably indicates that nitrate was not an important factor in this experiment. Nitrate may become toxic, however, in closed systems designed for very long periods between water changes. Wheaton (1977) states that most soluble phosphate in aquatic systems is in the form of inorganic orthophosphate. It is excreted by culture animals and also results from autolysis and subsequent mineralization of damaged or dead cells by heterotrophic bacteria (Spotte 1979). Goldblatt et al. (1978) also found small amounts leaching from gluten-based diets similar to the ration used in this experiment. Orthophosphate is removed from aquatic sys- tems by marine algae and by air stripping when the compound is absorbed onto the sur- face of air bubbles that rise to the surface. Results indicate that concentrations do in- crease in closed systems. Spotte (1979) states, however, that levels eventually reach equilib- rium because of air stripping. Toxic concen- trations have not been reported for lobster culture, but increased levels will stimulate al- gae growth in marine systems (Wheaton 1977). This algae may act as a dietary supple- ment for culture animals and may account for some of the increased growth in systems with low water exchange. Organic carbon results from animal wastes and extracellular products of aquatic plants (Spotte 1979). TOG levels may also include water soluble nutrients (vitamins, carbohy- drates, and proteins) that leach from unstable pelleted rations. TOG, like orthophosphate, is removed from water systems by air stripping (Spotte 1979). This may account for the ran- dom concentrations occurring between sys- tems. No toxic levels have been reported for lobster culture. All TOG levels reported were at least three times higher than those reported in large marine aquaria (Spotte 1979). This dis- crepancy occurred because of a difference in analysis procedure. Spotte used wet oxida- tion, whereas samples from this experiment were analyzed using dry combustion. Wil- liams (1975) reported three- to fourfold dif- ferences between these two techniques test- ing the same water sample. Mortality in most systems of both trials was very high. This was probably not a result of the water system, but occurred because of low nutritional value of the diet. High mor- tality has been an inherent problem in lobster culture with the use of formulated diets (Gonklin 1980). Recently, rations have been developed that consistently reduce mor- talities to less than 10 percent (Rex Infanger and Roger Mickelsen, unpubl. data). This is a significant advancement because low 9 8 * 7 E E I 6 i o o 5 z < 4 3 3 ' 2 1 ■ Fig. 4. Means and ranges of growth for trials 1 and 2. ° indicates significantly less growth compared with sys- tem V, using the standard Student t-test (p = .05). Fig. 5. Survival of juvenile lobsters from trials 1 and 2 after 90 days. 72 Great Basin Naturalist Vol. 42, No. 1 mortality and accelerated growth are neces- sary for lobster culture to be feasible. Closed systems provide many advantages over semiopen systems, including lower energy costs in heating and maintaining wa- ter temperature, greater efficiency in main- taining ideal culture conditions (salinity, pH, and dissolved oxygen), and fewer disease problems (King 1973). It can be concluded from this experiment that the closed systems tested also produce better growth of juvenile lobsters. Literature Cited Castell, J. D., AND J. F. Covey. 1976. Dietary lipid re- quirements of adult lobsters, Homarus american- iis (M.E.). J. Nutrition 106:1159-1165. Cobb, J. S. 1976. The American lobster: the biology of Homarus americanus. Univ. of Rhode Island Sea Grant Marine Tech. Rep. No. 49. 32 pp. CoNKLiN, D. E. 1980. Nutrition. Pages 277-300 in J. S. Cobb and B. F. Phillips, eds., The biology and management of lobsters. Academic Press, New York. Vol. 2. CoNKLiN, D. E., K. Devers, and R. a. Shleaser. 1975. Initial develpment of artificial diets for the lob- ster, Homarus americanus. Proc. World Mari Soc. 6:237-248. Dow, R. L. 1980. The clawed lobster fisheries. Pages 265-316 in J. S. Cobb and B. F. Phillips, eds.. The biology and management of lobsters. Aca- demic Press, New York. Vol. 2. Gallagher, M. L., R. C. Bayer, D. F. Leavitt, and J. H. Rittenburg. 1979. Formulation of artificial diets for feeding lobster {Homarus americanus) held in pounds. Maine Sea Grant Tech. Rep. No 46. Glude, J, B. 1977. NOAA aquaculture plan. U.S. Na- tional Oceanographic and Atmospheric Adminis- tration, Washington D.C. Goldblatt, M. J., D. E. Conklin, and W. D. Brown. 1978. Nutrient leaching from pelleted rations. Pages 117-129 in J. E. Halver and K. Tiews, eds., Finfish nutrition and finfeed technology. Heene- mann Verlags-gesellschaft, Berlin. Vol, 2. Infanger, R. C, R. W. Mickelsen, R. Heckmann, and S. R. Wadley. 1980. Vitamin leaching in lobster rations. Pages 3-10 in R. C. Bayer and A. D'Agostino, eds., Proc. Lobster Nutrition Work- shop, Maine Sea Grant Tech. Rep. No. 58. King, J. M. 1973. Recirculating system culture methods for marine organisms. SEA Scope 3:2,6-8. LiAO, P. B. and R. D. Mayo. 1972. Salmonid hatchery water reuse systems. Aquaculture 1:317-335 . Spotte, S. 1970. Fish and invertebrate culture, water management in closed systems. John Wiley and Sons, New York. 145 pp. 1973. Marine aquarium keeping. John Wiley and Sons, New York. 171 pp. 1979. Sea water aquariums. John Wiley and Sons, New York. 413 pp. Van Olst, J. C, J. M. Carlberg, and J. T. Hughes. 1980. Aquaculture. Pages 333-384 in J. S. Cobb and B. F. Phillips, eds.. The biology and manage- ment of lobsters. Academic Press, New York Vol 2. Wheaton, F. W. 1977. Aquacultural engineering. John Wiley and Sons, New York. 708 pp. Williams, P. J. 1975. Biological and chemical aspects of dissolved organic material in sea water. Pages 301-363 in J. P. Riley and G. Skirrow, eds.. Chemical oceanography. Academic Press, Lon- don. Vol. 2. Winget, R. R., R. E. Epifanio, T. Runnels, and P. Austin. 1976. Effects of diet and temperature on growth and mortality of the blue crab, Calli- nectes sapidus, maintained in a recirculating sys- tem. Proc. Natl. Shellfish Assoc. Vol. 66. 5 pp. DIAMETER-WEIGHT RELATIONSHIPS FOR JUNIPER FROM WET AND DRY SITES T. Weaver' and R. Lund' Abstract.- Height-diameter (basal or canopy) relationships for Jimipems scopulorum trees taken from wet and dry sites were quite different, but total aboveground weight-diameter relationships for trees taken from the two sites did not differ. It is shown that log total weight (kg) = approximately 1.7 -I- 2.26 log basal diameter (cm) = - 2.55 -I- 2.98 log canopy diameter (cm). Though the first relationship (i^ = 0.98) is stronger than the second (i^ = 0.80), the canopy diameter-weight relationship may be useful for estimating tree weights from aerial photos. Root-shoot ratios for wet site trees 5, 12, and 31 cm in basal diameter were 37, 27, and 26 percent, respectively. Linear dimensions are often well corre- lated with plant biomass (e.g., Kira and Shidei 1967, Weaver and Forcella 1977) and are therefore useful in estimating plant bio- mass. Basal diameter-weight relationships for many Rocky Mountain trees (Brown 1976, Weaver and Forcella 1977) and shrubs (Brown 1976) have been developed. Crown diameter-weight relationships for several Rocky Mountain shrubs are presented by Weaver (1977); this dimension may be espe- cially useful for low-multistemmed shrubs or for trees seen in aerial photographs. Because diameter-weight relationships are especially useful if they are general, it is de- sirable to test for generality by comparing di- ameter-biomass relationships of individuals of one species from different habitats and exhib- iting different life forms. Juniperus scopulo- rum Sarg. is a good test species because (1) it occupies both wet and dry sites, (2) it has a shorter and fuller form on dry than on wet sites, and (3) regression lines should be useful to managers throughout the Rocky Mountain area occupied by the tree (Little 1971). Methods Thirteen juniper trees were felled in Octo- ber 1980. They stood immediately east of the Headwaters of the Missouri State Park, Gal- latin County, Montana. Seven of these trees came from a bottomland site covered by a Fopulus trichocarpa T&cG— Juniperus scopulo- rum forest. The six remaining trees came from adjacent dry hills covered by a Juni- perus scopulorum-Bouteloua gracilis (HBK) Lag. ex Steud. woodland. The dry weight of each tree was deter- mined by the following procedure. The tree was felled and divided into portions with di- ameters of 0-1, 1-5, 5-10, and greater than 10 cm. These were weighed wet. Samples of material representative of each size were taken, weighed wet, and weighed again after drying to constant weight at 60 C. Dry weight/wet weight ratios for the 0-1, 1-5, 5-10, and 10-1- cm size classes were 61, 56, 61, and 61 percent on the dry site and 56, 54, 56, and 53 percent on the wet site. The dry weight of each tree was estimated by multi- plying wet weights of each portion by its dry weight/ wet weight ratio and summing across portions. The contribution of photosynthetic organs to the 0-1 cm class was estimated by sepa- rating green leaves and twigs from dried sam- ples, weighing, and expressing as a percent- age of the 0-1 cm total. The density of dry trunk wood was estimated by dividing sample weights by volumes (about 50 cc) esti- mated by displacement of water. The method used to test for the equality of regression lines in both wet and dry sites was to complete three separate regressions using data from, (1) wet sites only, (2) dry sites 'Department of Botany, Montana State University, Bozeman, Montana 59717. 'Department of Statistics, Montana State University, Bozeman. Montana 59717. 73 74 Great Basin Naturalist Vol. 42, No. 1 only, and (3) wet and dry sites combined. Us- ing the symbol SS, for the residual sum-of- squares in these regressions, the F-statistic for testing equal of regression lines in the two kinds of sites is F = (SS3 - SSi - SS2) (SSi + SS2)/(Ni + N2 - 4) ■ This value is compared to entries in an ordi- nary F table at 2 and Ni -f- Ng - 4 degrees of freedom. Although this procedure does not appear in statistical methods texts, it is a simple application of the basic theory for testing linear models (Graybill 1976). The new procedure is a reformulation of the anal- ysis of covariance problem to handle simulta- neously a test for both equality of slopes and intercepts (Snedecor 1980). Results and Conclusions Although the general climate of our two study sites was nearly identical, their envi- ronments were quite different. The climate of the Headwaters of the Missouri areas is dry-continental with an average annual pre- cipitation of 299 mm, average January tem- peratures of -1 C maximum to -12 C mini- mum, and average July temperatures of 28 C maximum to 11 C minimum (USDC 1978, Trident). The dry site trees came from a Bou- teloiia gracilis - Juniperiis scopidorum sav- annah typical of upland sites of the region. The wet site trees came from a riverside site dominated by old Populus trichocarpa trees; they undoubtedly experienced less water stress than the dry site trees due to the site's high water table and shelter from wind and radiation provided by the cottonwoods. lOi- 10 20 30 Basal Diameter (cm) ~ / - / Wei - ^ / A^ ^C - /^ -"' ^■^ Dry / ^ -A 3^" /-' 1 1 1 I 2 3 Crown Diameter (m) Fig. 1. Wet (A) and dry site (O) tree height-diameter relationships for both basal and canopy diameters differ significantly. Trees from different environments are ex- pected to have different forms, and indeed they do. (1) Though crown diameter at both sites is linearly related to tree height, at a given crown diameter, dry site trees are only half as tall as wet site trees (Fig. 1). We therefore describe dry site trees as 'broadly conical to hemispheric' and wet site trees as 'spindly.' (2) At both sites supporting struc- ture (basal diameter) increases logarith- mically with tree height as is mechanically necessary (Alexander 1971). Still, at a given height, trees of dry sites have bases almost twice as thick as those of wet sites. (3) Be- cause short trees of dry sites and taller trees of wet sites have similar quantities of small leafy twigs per unit of canopy breadth or bas- al diameter (Table 1), the trees of dry sites have denser foliage (more twigs per vertical meter) than the trees of wet sites. Despite larger reddish heartwood deposits, the den- sity of dry site trunk wood (0.49 ± 0.06 SE g/cc) is not significantly greater than that of wet site wood (0.42 ± 0.05 SE g/cc n = 3, p = 0.05). Despite the form differences just described, trees of both sites have essentially identical biomasses per unit of basal diameter or cano- py diameter. This is demonstrated visually with plots of untransformed total biomass data against tree size (Fig. 2). Regressions of log total aboveground weight (kg) and log 100 50 I 9 I I Total O -^^^' — / - / A ' - 9' ' To fa/ 1 / 1 1 1 1 / • / ' Twig 1 1 1 1 1 0 10 20 30 Basal Diameter (cm) 0 12 3 4 5 Crown Diameter (m) Fig. 2. Aboveground weight-diameter relationships are similar for trees grown on wet (A) and dry (O) sites. Dashed line summarizes total aboveground weight data and the solid line summarizes twig (0-1 cm diameter) weight data. March 1982 Weaver, Lund: Juniper Comparisons 75 Table 1. The relationship between tree diameter and aboveground tree weight is shown with regressions of the form log weight (kg or g) = a + b log diameter (cm). Site Wet Dry Pooled Wet Dry Pooled Wet Dry Pooled Wet Dry Pooled Intercept (a) Slope (b) Total weight (kg) basal diameter (cm) 1.78 2.21 0.98 1.55 2.37 0.98 1.70 2.26 0.98 Twig weight (g) basal diameter (cm) 1.64 1.78 0,98 1.60 1.98 0.98 1.59 1.91 0.97 Total weight (kg) canopy diameter (cm) -3.56 3.46 ^ ' 0.80 -1.94 2.70 0.98 -2.55 2.98 0.88 Twig weight (g) canopy diameter (cm) -2.59 2.75 0.79 -1.32 2.26 0.97 -1.93 2.48 0.85 7 6 13 7 6 13 7 5 12 7 5 12 0.72 2.67 0.54 1.19 'F testing for difference between wet and dry site regressions. Critical points are 95% = 4.26, 90% ■ 3.01, 75% = 1.62. twig biomass (g) against tree size (cm) are summarized in Table 1; in no case did popu- lations differ significantly. Separation of scales from leafy twigs shows that 75 ± 3 and 59 ± 6 SE percent of the leafy twig weight is leaf on dry and wet sites respective- ly; with six samples from each tree the differ- ence is not statistically significant at p = 0.05. Root-shoot ratios for bottomland trees 5, 12, and 31 cm in diameter were 37, 27, and 26 percent, respectively. Root-shoot ratios appear to generally decline with in- creasing tree size (Weaver 1974). Our observation that tall trees and short trees of the same diameter have the same weight is both counter-intuitive and useful. Even if dry site wood is slightly more dense than wet side wood, density differences are insufficient to explain this observation. In- stead, the biomass appears to be distributed as if there were a fixed amount of photo- synthate per internode which is devoted to diameter growth if it is not expended in height growth. Such allocation on the dry site could be due to adequate light combined with trunk thickening induced by wind stress, while on the wet site it might be due to shade induced etiolation combined with low wind stress (Kramer and Kozlowski 1979, Zimmerman and Brown 1971). Regardless of its cause, the relationship is useful because it suggests that we may apply diameter-biomass relationships developed on one site to other, environmentally dissimilar sites. Acknowledgments We gratefully acknowledge the help of K. Boggs, J. Clarke, B. Goulding, S. Gregory, C. Hunt, J. Lichthardt, D. MuUan, J. Newbauer, and K. Striby in harvesting the trees. Literature Cited Alexander, R. 1971. Size and shape. Edward Arnold, London. 59 pp. Brown, J. 1976a. Predicting crown weights for eleyen Rocky Mountain conifers. Pages 103-115 in Oslo Biomass Studies. International Union of Forest Research Organizations (lUFRO) Congress. 1976b. Estimating shrub biomass from basal stem diameters. Canadian J. Forest Research 6:15.3-158. Grayhill, F. 1976. Theory and application of the linear model. Duxbury Press, Belmont, California. KiRA, T., AND T. Shidei. 1967. Primary production and turnoyer of organic matter in different forest eco- systems of the western Pacific. Jap. J. Ecology 17:70-87. Kramer, P., and T, Kozlowski. 1979. Physiology of woody plants. Academic Press, New York. 811 pp. Little, E. 1971. Atlas of United States trees. Vol. I: con- ifers and important hardwoods, USDA Forest Seryice Misc, Publ, 1146. U.S. Goyt. Printing Off., Washington, D.C. 210 pp. Snedecor, G., and W. Cochran. 1980. Statistical meth- ods. Iowa State Uniy. Press, Ames, Iowa. 76 Great Basin Naturalist Vol. 42, No. 1 Weaver, T. 1974. The growth of two scrub oaks and the cost of their leaves. Montana Acad. Sci. Proc. 34:19-23. 1977. Area-mass relationships for common Mon- tana shrubs. Proc. Mont. Acad, of Sci. 37:54-58. Weaver, T., and F. Forcella. 1977. Biomass of fifty conifer forests and nutrient exports associated with their harvest. Great Basin Nat. 37:395-401. U.S. Department of Commerce. 1978. Climatological data. National Oceanographic and Atmospheric Administration (NOAA) Environmental Data Ser- vice, National Climatic Center, Asheville, North Carolina. Zimmerman, M., and C. Brown. 1971. Tree structure and function. Springer Verlag, New York. 336 pp. A DESCRIPTION OF TIMPIE SPRINGS, UTAH, WITH A PRELIMINARY SURVEY OF THE AQUATIC MACRORIOTA Thomas M. Baugh', Michael A. Nelson', and Floyd Simpson' Abstract.— a description of some physical, chemical, and biotic features of Timpie Springs, Utah. Timpie Springs, Tooele County, Utah, is a spring, pond, marsh complex (Fig. 1) in the southeastern comer of the intersection of In- terstate 80 and the Skull Valley Road (Utah Highway 40), Sections 8 and 9, Range 7 W, Township IS. This aquatic complex is at an altitude of 1300 m at the northwestern base of the Stans- bury Mountains. With the exception of the spring and related mesic habitat, the sur- rounding area is a typical, xeric, cold-shrub Great Basin sit. Greasewood (Sarcobatus ver- miculatus), samphire {Salicomia sp.), and goat grass {Aegilops cylindrica) are the domi- nant riparian vegetational species. Present human use of the area includes cattle watering and grazing, fishing, hunting, water withdrawal for road and other con- struction projects, and water for the im- poimdments of the adjacent Timpie Wildlife Management Unit. Human modifications of the area include a galvanized steel flume in the channel con- necting the pond with the marsh, a levee sep- arating the marsh from the pond, and a double culvert draining the marsh waters un- der Interstate 80 into the Timpie Wildlife Management Unit. A well-graded gravel road leads from the Skull Valley Road west to the spring where it joins another well-graded gravel road that follows the foot of the Stans- bury Mountain Range in a roughly north- south direction. Timpie Springs lies at the extreme north- western tip of the north-south-trending Stans- bury Mountain Range, the northern portion of which is characterized as the Garden City Fig. 1. Schematic of Timpie Springs, Tooele Co. Utah. '1020 Custer Avenue. Ogden, Utah 84404. '3765 Harnson Boulevard, Ogden, Utah 84403. 77 78 Great Basin Naturalist Vol. 42, No. 1 Formation (Rigby 1958). The spring issues from the easternmost of a series of faults in the Stansbury Range (Everett 1957). Accord- ing to Everett (1957) all groundwater in Skull Valley is produced from deposits of lake clays, silts, and gravels intermixed with allu- vial deposits. Everett (1957) divides the groundwaters of the valley into a northern saline reservoir and a southern freshwater reservoir. Timpie Springs, originating in the northern reservoir, is estimated to contribute 20,000 tons of salt (total salt in solution) an- nually to the Great Salt Lake (Eardley 1957). Materials and Methods Water chemistry values were measured on three occasions at three points in the system: Point A = spring effluent, B = channel, and C = marsh. Chemical constituents were measured by the Hach' drop count method and converted to mg/1, where applicable. Temperature was measured with a Weston thermometer (Model 2265). Fish were captured with long-handled dip- nets from within six feet of the banks of the pond and marsh and throughout the length of the channel. The presence or absence and relative abundance of each species was noted. Aquatic macroinvertebrates were sampled nonquantitatively along the banks of the pond and marsh and throughout the channel. The presence or absence and relative abun- dance of a species was noted. Aquatic macroflora was sampled non- quantitatively along the banks of the pond and marsh and throughout the channel. The presence or absence and relative abundance was noted. Results Pond.- Dabb (1977, 1980) reports the mean depth of the Timpie Springs pond as 5 ft. Our measurements give a surface area of 117,500 ft^ for the pond. To reach an esti- mate of total volume, however, it appeared necessary to apply the 5 ft depth only to the main body of the pond. Inclusion of the two main areas of shallows (Fig. 1) gives an inflat- ed figure for volume. The shallow areas ac- count for 34,500 ft^ of surface area but, at an average depth of one foot, only 34,500 ft^ of volume. This leaves a surface area of 83,000 ft^ for the main body of the pond or a volume for the main body of 415,000 ft^, bringing the total pond volume to 449,500 ft^. The pond substrate consists mostly of a deep deposit of silt and sand. Channel.— Discharge from the pond into the channel is 4.6 cfs. Discharge (Q) was cal- culated by the method described by Reid and Cox (1976): Q(cfs) = WD„V„, where Q = discharge (cfs) Dn, = mean channel depth W = width (ft) Vn, = mean velocity (ft/sec) 'Hach Chemical Company, Loveland, Colorado. Table 1. Temperature and chemical constituents, Timpie Springs, Tooele Co., Utah. Site and date in Temperature (C) pH TH' TA^ Ca Mg CI DO' October 1981 mg/1 A. 3 18.0 7.5 721.5 189.0 395.1 481.0 481.0 5 17 17.8 7.5 652.8 189.0 326.4 326.4 439.5 5 31 17.8 7.5 670.0 189.0 360.7 309.4 451.5 5 B. 3 17.8 8.0 721.5 206.1 377.9 343.6 535.0 8 17 14.4 8.0 670.5 189.0 343.6 326.4 478.8 6 31 14.4 8.0 652.8 189.0 377.9 274.9 _ 7 C. 3 16.4 8.0 755.9 171.8 412.3 343.6 530.0 6 17 13.2 8.0 738.7 171.8 395.1 343.6 500.0 7 31 6.1 8.0 687.1 189.0 377.9 309.2 463.6 8 'Total hardness 'Total alkalmitv 'Dissolved oxygen March 1982 Baugh et al.: Timpie Springs 79 The channel substrate is a loose aggregate of silt and sand above the flume and gravel and rock below the flume to the confluence of the channel with the marsh. Marsh.— The marsh is a shallow feature with a surface area of 145,945 ft^, mean depth of about one ft, and a total approx- imate volume of 145,945 ft^. It should be noted that marsh depth can vary as much as 50 percent, depending on water use at the Timpie Wildlife Management Unit. The marsh substrate is a deep, loose aggregate of silt, sand, and detritus. Temperature and chemical con- stituents.— Temperature and the chemical constituents of Timpie Springs are reported in Table 1. It should be noted that Ave days of heavy rain preceded the measurements taken on 17 January 1981. Fish.— Four species of fish Gila atraria (Utah chub), Lucania parva (rainwater kill- ifish), Gamhusia affinis (mosquitofish), and Micropterus salmoides (largemouth bass), oc- cupy one or more of the components of the Timpie Springs system (Table 2). According to Sigler and Miller (1963), M. salmoides and L. parva were introduced prior to 1959, M. salmoides to develop a sport fishery, and L. parva accidentally. It is probable that G. af- finis was introduced to control mosquitoes. All populations of all species are self- sustaining. Aquatic macroflora.— The major species of aquatic macroflora are listed in Table 3. Aquatic macroinvertebrates.— The ma- jor aquatic macroinvertebrates of Timpie Table 2. Fishes of Timpie Springs, Tooele Co., Utah. Species Location' Frequency- Gila atraria Gamhusia affinis Lucania parva Micropterus salmoides P C M P C M P C M P C M C C C VC C VC C C VC C NP NP Springs are listed in Table 4. It should be noted that both the occurrence and numbers of many invertebrates are dependent on sea- sonal climatic conditions. For example, the week prior to our sampling of 17 October 1981 was cool, with air temperatures ranging from 1.6-4.4 C. Both larval and adult mos- quitoes, which had been numerous on 3 Oc- tober 1981, were conspicuously and pleas- antly absent on 17 October 1981. Summary Timpie Springs is an aquatic system sur- rounded by typical cold-shrub desert. Major human modification includes the in- troduction of three species of nonnative fish, the construction of gravel roads, and grazing by cattle. Recreation use includes fishing and some hunting. The spring waters are neces- Table 3. Major aquatic macroflora of Timpie Springs, Tooele Co., Utah. Type and genus Location' Frequency' Pepperwort Marsilea sp. Water nymph Najas sp. Blue-green algae Enteromorpha sp. Oscillatoria sp. Lijngbya sp. Green algae Chara sp. Cladophora sp. Spriogyra sp. P C M P C M P C M P C M P C M P C M P C M P C M C NP R VC VC VC VC C VC C R C VC R UC C R VC UC R UC UC R UC 'P = pond, C = channel, M = marsh. 'VC = very common, C = common, UC = uncommon. R = rare, NP not present. 'P = pond, C = channel, M = marsh. 'VC = very common, C = common, UC not present. uncommon, R = rare, NP = 80 Great Basin Naturalist Vol. 42, No. 1 Table 4. Major aquatic macroinvertebrates of Timpie Springs, Tooele Co., Utah. Type Order Location i' Frequency- Mosquito Diptera C M P VC vc vc Dragonfly Damselfly Odonata P C M c c c Mayfly Ephemeroptera P C M c c c Backswimmer Hemiptera P C M vc c vc Crustacean Amphipoda (Hyalella) P C M vc vc vc Snail (Gastrapoda) P C M vc vc vc 'P = pond, C = channel, M = marsh. 'VC = very common, C = common, UC = uncommon, R = rare. sary to maintain the aquatic habitats of the closely adjacent Timpie Wildlife Manage- ment Unit. Acknowledgments We thank Dr. Earl Smart, Department of Zoology, Weber State College, and Donald Andriano, Jim Johnson, and Glenn Davis, Utah State Division of Wildlife Resources, for their reviews and helpful comments. We thank Dr. Eugene Bosniak, Department of Botany, Weber State College, for his help in identifying the aquatic macroflora. Literature Cited Dabb, B. 1977. Work study sheets for lakes, reservoirs, and ponds. Report of the Utah State Division of Wildlife Resources, Salt Lake City. 1980. Work study sheets for lakes, reservoirs, and ponds. Report of the Utah State Division of Wildlife Resources, Salt Lake City. EaRDLEY, a. J., V. GVOZDETSKY, AND R. E. MaRSELL. 1957. Hydrology of Lake Bonneville and sedi- ments and soils of its basin. Geol. Soc. Amer. 68:1141-1202. Everett, K. R. 1957. Geology and groundwater of Skull Valley, Tooele County, Utah. Unpublished thesis. Univ. of Utah. 92 pp. Reid, G. K., and R. D. Wood.. 1976. Ecology of inland waters and estuaries. D. Van Nos tand Company, New York. 485 pp. RiGBY, J. K. 1958. Guidebook to the geology of Utah, No. 1.3, Geology of the Stansbury Mountains, Tooele County, Utah. Utah Geological Soc. 175 pp. Sigler, W. F., and R. R. Miller. 1963. Fishes of Utah. Utah State Department of Wildlife Resources, Salt Lake City. 203 pp. VEGETATION AND SOIL FACTORS IN RELATION TO SLOPE POSITION ON FOOTHILL KNOLLS IN THE UINTA BASIN OF UTAH Miles O. Moretti' and Jack D. Brothersoir .\bstract.- Vegetation and soil differences with respect to slope position were studied on foothill knolls in the Uinta Basin of Utah. Plant communities on windswept ridges (top of slope) exhibited several unique characteristics when compared to the other slope communities. These communities at the top and base of slopes were sufficiently different in respect to plant life form composition, plant cover, wind-adapted growth forms, and percent exposed rock that they should be considered separate community types. Mineral concentrations in plant tissue and soil sam- ples declined downslope in some cases and increased in others. Diversity decreased downslope as shrubs became dominant over grasses and forbs. Management of these communities should require special consideration due to the changes in the community structure with slope position. Vegetation composition, soil factors, and tlieir changes in response to exposure and slope position have received increased atten- tion in recent years (Harner and Harper 1973, Jaynes and Harper 1978, Bloss and Brotherson 1979). The foothill knolls in the Uinta Mountains of Utah are little known ecologically and thus offer opportunities to further our knowledge of such relationships. These foothill knolls, many of which lie adja- cent to mountain stream drainages, receive incessant winds, thus creating a unique plant community. Ecological studies of windswept plant communities have generally been re- stricted to high alpine ridges (Marinos 1978). Little research has been done on such com- mmiities at lower elevations (i.e., 2000 and 3000 m). Anderson et al. (1976) described wind- swept ridges in south central Wyoming as a vegetative type. The areas so described were ridges that received strong southwesterly winds and had a cover of mat-forming plants. Establishment of the sagebrush-grass commu- nity of less win^y spots in the area was ap- parently precluded l3y the winds. The domi- nant plant species was Lyal's goldenweed (Haplopappus hjallii). Plant cover for the community was estimated to be about 33 percent. Soils in the area were moderately textured (sandy loam to sandy clay loam), with soil depth averaging 17 cm. Exposed rock was twice as great at the top of the slope as at the bottom. Mineral concentrations in vegetation and soil with respect to slope have not been stud- ied to any extent. Harner and Harper (1973) looked at the increase in mineral concentra- tion in vegetation along a moisture gradient. They concluded that increasing soil moisture permits greater solubility and, therefore, greater absorption of minerals in more pro- ductive sites. Fairchild and Brotherson (1980) used topographic position and slope aspect data as independent variables in statistical analyses of shrub habitats in conjunction with mineral concentration in the shrubs and soil. The objective of this study was to deter- mine the ecological relationships (vegetative and soil) of plant communities, some of which are windswept, with respect to slope position. Knowledge of vegetation and soil differences with respect to slope position should be useful for range managers planning treatment programs (i.e., brush control, range reseeding, etc.). Such would be especially true in areas similar to those described in this study, because much of Utah's big game win- ter ranges occur in similar topographic locations. 'Division of Wildlife Resources-State of Utah, 455 West Railroad Avenue, Pnce, Utah 84501. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 81 82 Great Basin Naturalist Vol. 42, No. 1 Study Area The Uinta Basin is characterized as a broad, elongated, asymmetric basin lying in the northeastern corner of Utah and extend- ing into northwestern Colorado (Brotherson 1979). The central portion of the Uinta Basin is considered a cold desert giving rise to ex- tensive pinyon-juniper forests with a book- cliff type topography (Greenwood and Broth- erson 1978). The foothill areas of the Uinta Mountains that lie above the pinyon-juniper forests are dominated by mountain brush. The study area (north and west of Mountain Home, Utah) was located in the mountain brush vegetational zone on three foothill knolls at the foot of the Uinta Mountains. The elevation of the area ranges between 2400 and 2800 m. Climate is highly variable, wdth cold winters and hot dry summers. An- nual precipitation in the area is between 30 and 35 cm, coming mostly in the form of snow during the winter. The area is used as late winter and early spring range for sheep and winter range for elk, deer, and sage grouse. The communities studied were similar in that all areas display northeast-southwest ex- posure. Winds generally blow out of the west and are channeled down mountain stream drainages to the west of the study areas. The windswept side of the communities, especially near the crest of the hill, is domi- nated by low mat-forming plants, some with the life form of a forb above ground and a shrub below ground. The leeward side is dominated by plants similar to those found on the windward side near the top. The vege- tation downslope, where soils are deeper and receive increased moisture from snowdrifts, is dominated by tall shrubs. Methods Three hills were studied within a 26 km2 area. All had similar exposure, slope, and ele- vation to insure that meaningful comparisons could be made. Each hill was sampled on both northeast and southwest sides at the top, middle, and bottom slope positions. A total of 90 plots (3 X 3 m) were sampled (30 plots per hill and 15 plots per side). Five plots each were placed along the contour at the top, middle, and bottom of each hill. Study areas were marked with nylon cord 12 m long with a loop tied every 3 m. The four corners were secured with wooden stakes. Flagging of alternate colors was tied at equal intervals to help insure uniform placement of the 0.25 m2 quadrats used to subsample each area. Four subsamples were taken within each 3 X 3 m area. Density of shrubs was determined from direct counts of all shrubs within the 0.25 m2 quadrats. Per- cent cover was taken in each 0.25 m2 quadrat for each plant species, rock, bare ground, and litter. Cover values were estimated as sug- gested by Daubenmire (1959), with some modification. The cover classes: (1)0.01-1.0 percent, (2)1.1-5.0 percent, (3)5.1-25 per- cent, (4)25.1-50 percent, (5)50.1-75 percent, (6)75.1-95 percent, and (7)95.1-100 percent were weighted to the lower end, in an at- tempt to make the system more sensitive to large numbers of low-growing species of plants and not to overestimate those plants near the top of the ridges where percentage of exposed rock is greatest. Height of species was also measured. Where a species was rep- resented by less than 5 individuals per plot, all individuals were measured. For species represented by more than 5 individuals per 0.25 m2 quadrat, 5 individuals were selected at random for measurement. Soil penetration was measured with a 1-m penetrometer. Depth measures were taken 5 times in each 0.25 m2 plot. Measurements were taken once at each corner and once in the middle. A total of 20 soil depth measure- ments was taken within each 3 X 3 m plot. Soil samples were taken from each comer and middle of each 3 X 3 m plot. These 5 samples were then combined to get a repre- sentative sample of the whole plot. Soil sam- ples were analyzed for texture, pH, soluble salts, and mineral ion concentrations (i.e., cal- cium, magnesium, potassium, sodium, iron, manganese, zinc, copper, nitrogen and phos- phorus). Texture was determined as suggested by Bouyoucos (1951). Soil reaction was deter- mined with a glass electrode pH meter. So- luble salts and pH were determined on satu- rated soil pastes having a 1:1 soil to water ratio (Russell 1948). Organic matter was de- termined by heating soil samples for 24 hours March 1982 MoRETTi, Brotherson: Vegetation-Soil Factors 83 at 450 C. Differences in weight before and after heating were converted to percent or- ganic matter. Three soil samples from each slope posi- tion were randomly chosen out of the five taken for analysis of mineral concentrations. Individual ion concentration was determined by a Perkin-Elmer Model 403 atomic absorp- tion spectrophotometer (Isaac and Kerber 1971). Potassium, magnesium, calcium, and sodium ions were extracted with a 1 percent neutral normal ammonium acetate solution (Jackson 1958, Hess 1971, Jones 1973). Zinc, manganese, iron, and copper were extracted by using DTPA-diethylenetriaminepenta- acetic-acid extracting agent (Lindsay and Norvell 1969). Soil phosphorus was extracted by sodium bicarbonate (Olsen et al. 1954). Total nitrogen analysis was made using a macro-kjeldahl procedure (Jackson 1958). Plant samples for chemical analysis were taken randomly at each slope position from within the 3 X 3 m plots. Current year growth for shrubs and all aboveground tissue for forbs and grasses was clipped. The sam- ples were air dried and ashed at 450 C for 24 hours. The ash analysis was made with the atomic absorption spectrophotometer. Results and Discussion The environmental factors measured in this study to characterize the communities on foothill knolls near the Uinta Mountains ex- hibited several general patterns (Table 1). Soil depth was found to be greatest in the midslope position of both the windward and leeward sides (18.5 and 27.7 cm respectively) of the hills. This is possibly the result of suf- ficient moisture to move the soil from the tops of the ridges but not enough to force the soil outward from the slope base. The lee- ward position is unique with respect to the other positions as the area receives most of its moisture from the increased accumulation of snow from drifts due to snow accumulation on the leeward side. The areas also have a northeasterly exposure; thus their snow cover remains longer. The result is increased mois- ture being available to plants for longer peri- ods during critical early growth stages. The leeward position was dominated by Big Sage- brush {Artemisia tridentata) and Utah Ser- viceberry {Amelanchier utahensis). The highest percent exposed rock (61.9 percent) was found on the windward side at the top-slope position; rock then steadily de- creased downslope (Table 1). Top positions receive the greatest impact from winds, which sweep away snow and much soil and leave exposed rock. The leeward position shows much less exposed rock (25 percent) but displays a similar trend downslope. Mid- slope positions probably receive part of the soil blowing in from the windward side. Ex- posed bare ground and litter cover generally increased downslope. Organic matter and to- tal soluble salts increased slightly downslope (Table 1), and pH showed slight decreases. Plant life form data (Table 2) give an in- dication of patterns .produced by variation in the relative cover of each life form class. Shrubs increased in relative cover downslope on both sides of the knolls. In contrast, shrub density decreased slightly downslope. This was due to larger plants at the bottom (Table 2) giving a lower density of shrubs but a greater percentage of cover. The midslope position had more species of small shrubs than did the bottom. The shrubs in the top position were small and mat forming, thus giving the lowest cover value for the shrubs (6.0 percent). Table 1. General environmental factors downslope on windswept communities. Environmental factor Slope position windward side Slope position leeward side Top Middle Bottom Top Middle Bottom Soil penetration Bare ground (%) Rock (%) Litter (%) Organic matter (%) pH Soluble salts (ppm) 6.7 20.1 6L9 27.3 6.6 7.1 181 18.5 40.0 38.3 30.5 8.1 7.0 432 16.4 41.0 13.2 46.0 8.3 6.9 357 6.7 20.8 25.2 41.8 7.4 6.9 207 27.7 14.9 4.0 67.7 6.6 6.9 162 21.6 21.1 14.7 59.4 6.9 6.7 275 84 Great Basin Naturalist Vol. 42, No. 1 Forbs showed their highest cover (13.6 percent) at the top of the windward side of the slope (Table 2) and then decreased down- hill. The windward side decrease was much greater than the leeward side. Forbs were poorly represented on the leeward side. The increased density and percent shrub cover on the leeward side probably crowd out forbs. Grasses showed a relatively constant per- cent cover on both sides (Table 2). The wind- ward side had a slight decrease in percent cover, but the leeward side had a slight in- crease. Both had lower percent cover values at the midslope position than at the other two. At the top of the windward side, grasses had a greater percent cover (14.8 percent) than any of the other life form types. Percent cover for plant species thought to be successful on the windswept ridges (i.e., showed growth forms adapted to windy con- ditions) were considered together in a group called wind-adapted plants. Plants in this group were mat forming, rhizomatous, or formed rosettes. Cover for species in this group (15.6 percent) was greatest at the top of the windward side (Table 2). The top posi- tion of the leeward side had the second larg- est cover value (8.3 percent). The lowest cov- er value (1.1 percent) for this group was at the midslope position of the leeward side, which was dominated by tall shrubs. Height of plants was greatest, soil penetration was greatest, and moisture is assumed to be max- imal because the largest snowdrifts accumu- lated there. The Uinta Basin experienced a drought throughout the first half of 1977, the spring being extremely dry following record low moisture accumulations during the winter. This may explain the low percentage of an- nuals in the study area. The annuals either did not germinate that year or appeared be- fore the study was initiated. The average height of all vegetation at each slope position closely paralleled soil depth on the same slope. Vegetation height and soil depth were significantly (positive) correlated (P < 0.01), both tending to in- crease downslope on both windward and lee- ward sides (Table 2). The leeward side, how- ever, showed an increase in height at the middle position (24.6 cm), then a decrease again at the bottom (19.2 cm). The increase in height can be accounted for by the pres- ence of populations of Utah serviceberry at this midslope position. A diversity index (Levins 1966, MacArthur 1972) was determined by using percent cover of each plant at each position (Table 2). Di- versity decreased on both sides of the knoll downslope. The top position on the leeward side had the highest diversity (9.4). The low- est diversity (3.5) was at the bottom of the leeward side. A Shannon-Weiner (1949) di- versity index was also computed and showed similar trends to that of the 1/Sum Pi2 diver- sity index. Cluster analysis (Sneath and Sokol 1973) was used to group slope positions on the basis of vegetative cover (Fig. 1). The bottom- slope positions of both sides of the knolls were the most similar with respect to each other. The top-slope position on the wind- ward side of the knolls was the most different vegetatively of any of the slope positions with respect to all others. This and other evi- dence indicates that the top of the slope on the windward side is unique with respect to Table 2. General biotic factors downslope on windswept communities. Biotic Slope position windward side Slope position leeward side factor Top Middle Bottom Top Middle Bottom Life form (% cover) Shrub 6.0 21.0 21.4 19.3 26.7 30.0 Forb 13.6 2.0 4.4 4.8 4.4 2.9 Grass 14.8 9.2 12.0 14.6 13.5 15.1 Annual 0.2 0 0.1 0 0.2 0.3 Wind-adapted plants 15.6 .3.6 3.7 8.3 1.1 2.6 Shrub density 8.7 7.3 6.0 11.9 12.1 8.9 X height (cm) 9.4 16.7 17.1 12.0 24.6 19.2 Diversity (1/ Sum F^) 6.9 4.4 3.9 9.4 6.4 3.5 X No. of plant species 14.7 10.3 13.0 17.0 17.7 16.7 March 1982 MoRETTi, Brotherson: Vegetation-Soil Factors 85 30 .JL- % Similarity e.0 60 80 90 100 Windward-Top Leeward-Top Leeward-Midslope Windward-Midslope Windward-Bottom Slope Leeward-Bottom Slope Fig. 1. Phenogram of the relationships between the slope position communities. Cluster is based on species cover values. the remainder of the hill. There exists an area that could well be classified as a separate vegetation type with regard to the surround- ing plant community. Anderson et al. (1976) stated that windswept communities are re- stricted to the ridges where the more com- petitive sagebrush can't tolerate the desiccat- ing winds and rockier soil. The areas provided habitat for low, mat-forming plants. The area had an average plant cover of 33 percent and compares closely to this study, which has an average plant cover in the same position of 34.6 percent. The average number of plants per slope position used to develop a prevalent species list is shown in Table 3. The windward side Table .3. Prevalent species (% cover) for foothill knolls in the Uinta Basin, Utah. Species Artemisia tridentata Xanthocephahim sarothrae AmelancJiier utahensis Phlox bnjoides Koeleria cristata Stipa coinata Antennaria parvifolia Boiiteloua gracilis Poa fendleriana Artemisia cana Artemisia frigida Agropyron spicatiim Eriogonum heracleoides Berberis repens Penstemon sp. Symphoricarpos orcophilus Ehjmiis ambiguus Purshia tridentata Cercocarpus montanus Chrysothammts greenei Slope position windward side Top Middle Bottom 0.0 0.4 0.0 10.3 1.4 4.6 0.0 1.6 0.0 0.8 4.6 4.4 0.0 0.0 0.0 0.0 1.8 0.0 0.0 0.0 1.3.6 0.4 0.4 1.7 0.4 0.0 0.0 1.1 1.0 0.2 1.1 4.8 0.0 0.0 0.0 0.0 1.8 0.0 4.9 0.0 18.3 0.0 0.4 3.4 2.2 1.0 0.2 0.0 1.1 0.1 0.0 6.1 0.1 0.0 0.0 1.7 1.5 0.0 0.0 0.4 Slope position leeward side Top Middle Bottom 11.5 2.2 2.1 0.4 0.1 2.6 0.3 4.1 4.0 0.8 0.6 1.7 1.0 0.9 0.6 0.6 0.0 0.0 0.0 0.0 11.8 0.0 5.9 0.6 1.6 0.8 0.0 0.0 0.0 0.0 0.0 8.1 1.1 0.4 1.6 8.0 0.0 0.0 0.0 0.0 23.8 0.9 0.0 1.1 1.1 0.6 0.4 0.0 0.8 0.0 0.0 8.8 0.0 0.0 0.2 2.6 4.2 2.3 0.0 0.0 86 Great Basin Naturalist Vol. 42, No. 1 had an average of 12.7, but the leeward side was much higher with 17.1 species. A prevalent species list was developed for each slope position based on the average number of species per position and percent- age cover (Table 3). Plant cover was general- ly low at most positions. Forbs and grasses exhibited reduced growth due to soil droughtiness, thus contributing less to the overall percent cover. Total percent cover increased downslope. The lowest percent plant cover (37.0 percent) on the leeward side was higher than the highest percent cov- er value (36.7 percent) on the windward side. The largest percent plant cover (46.4 per- cent) was at the bottom position of the lee- ward side of the knoll but had the highest concentration of a single species to the total. Big sagebrush (Artemisia tridentata) provided 23.8 percent cover (51 percent of the total plant cover for the bottom position). The highest percent cover on the windward side was again at the bottom position (36.7 per- cent), where big sagebrush (18.3 percent ab- solute cover) contributed 50 percent of the total cover (Table 3). Two species were found in study plots at all slope positions: Moss phlox (Phlox bnjoides) and Bluebunch wheatgrass (Agropij- ron spicatum). Big sagebrush exhibited the highest percent cover of any species at all ex- cept on the top slope position on the wind- ward side, where it did not occur. The sites on the windward side were dominated by Moss phlox (10.3 percent); Needle and Thread (Stipa comata), (4.6 percent cover); Fringed sagebrush (Artemisia fngida), (4.6 percent); and Bluebunch wheatgrass (4.4 percent). Soil ion concentrations ranged from a low of 0.6 ppm for copper at the midslope posi- tion on the leeward side to a high of 5947 ppm for calcium at the midslope position on the windward side (Table 4). Concentrations of minerals between windward and leeward sides generally followed similar patterns. Zinc is the only ion to decrease downslope on one side and increase downslope on the other. Calcium and iron generally decreased downslope, but phosphorus, manganese, ni- trogen, magnesium, and potassium showed increases (Table 4). Nitrogen, calcium, mag- nesium, and potassium occurred in high con- centration on both exposures. Epstein (1972) suggested that for soils to maintain healthy plant tissue, a soil concentration of 200 ppm of phosphorus is required. Both exposures had 10 percent or less of this concentration of phosphorus in the soil. Mineral concentrations in plant tissue re- mained level or generally increased down- slope on the windward and leeward expo- sures. Ion concentration in the plants ranged from 8 ppm for copper on the top position of the leeward side to 23,000 ppm for nitrogen at the midslope position of the leeward side (Table 5). The high concentration of nitrogen in the midslope position on the leeward side corresponds with the highest percent litter (67.7 percent). This can be attributed to an increase in nitrogen fixation by free-living microorganisms in and under litter mats (Charley 1977). The mean ion concentration for copper in plant tissue was considerably lower (10 ppm) than the mean of 28.9 ppm found by Broth- ersen and Osayande (1980) in True Mountain Mahogany (Cercocarpus montanus) 30 km to Table 4. Soil mineral concentration downslope on windswept communities. Minerals (ppm) Slope position windward side Top Zinc Iron Phosphorus Manganese Copper Sodium Nitrogen Calcium Magnesium Potassium 1.3 16.4 11.9 12.0 0.8 42.0 2000. 3971. 168. 169. Middle 1.4 10.9 10.8 11.3 0.9 42.3 1400. 5947. 606. 232. Bottom 1.5 14.5 15.8 18.0 1.2 41.0 1800. 3790. 491. 300. Slope position leeward side Top 2.1 21.2 13.9 12.7 0.7 40.3 2100. 3378. 223. 208. Middle 1.6 21.1 13.8 15.0 0.6 .35.7 2000. 2853. 230. 200. Bottom 1.2 17.0 20.1 18.7 0.9 41.0 1900. 2558. .338. March 1982 MoRETTi, Brotherson: Vegetation-Soil Factors 87 the southeast in the pinyon-juniper wood- land. The concentration is well below the 20 ppm suggested by Baker (1974) as being toxic to some ruminant animals. The highest con- centrations of copper (11 ppm) were found at the bottom positions of both exposures. Concentrations of the macronutrients (iron, phosphorus, nitrogen, calcium, magnesium, and potassium) in the plants were high and the concentrations of micronutrients were low. Ion concentrations in the plants were found to be adequate for higher plants to maintain a healthy condition (Epstein 1972). Minerals concentrated in plant tissue varies in meeting the nutritional requirements of sheep and cattle that graze the area. Most minerals met or exceeded the mineral re- quirements of sheep and cattle (NRC 1976). Copper (8-11 ppm) and iron (308-706 ppm) were the only two elements present in quan- tities considered to be toxic. Copper within the 8-25 ppm range is considered to be toxic to sheep. Iron exceeded the toxic level for cattle (400 ppm) in three of the six slope po- sitions. Because the mineral concentration in plant tissue was determined from a com- bination of all the plants within each area, some plant species high in these minerals may be impalatable to animals and not part of their diet. The ratio of mineral concentration in the plant versus the soil shows to what extent the plants take up and concentrate the minerals (Table 6). The highest ratio was found for phosphorus, which was 108 to 176 times more concentrated in plant tissue than in the soil. The lowest ratio for mineral concentra- tion was for manganese (2.1) at the bottom- slope position of the leeward side. Manganese had the lowest concentration ratio for all the minerals at all of the positions (Table 6). Potassium had the second highest ratio, ranging from a low of 48.3 at the bottom- slope position to a high of 81.0 at the top, both on the windward side. Potassium, the only monovalent cation essential for all high- er plants, is inefficient as a cofactor in en- zyme systems and plants have evolved the ability to take up large concentrations from the soil (Epstein 1972). The mineral concentrations in the soil and plants, plant life form, and other environ- mental factors were subjected to correlation analysis (Table 7). Several significant correla- tions developed within and among the groups in relation to changes downslope. Soil depth was positively correlated with plant height; thus, as soil became deeper downslope, plant height increased. Soil depth was negatively correlated with plant density; thus, as shown earlier, shrub density was highest at the top of the hills where soil was shallowest (Table 7A). Shrubs as a life form were negatively cor- related with forbs. Consequently, as shrubs increased downslope forbs decreased (Table 7 A). Cover for forbs (13.6 percent) was high- est at the top position on the windward side, where shrub cover (6.0 percent) was lowest. Shrubs were also highly correlated with plant height. Forbs were negatively correlated with height downslope. Percent cover by forbs (13.6 percent) was highest at the top position on the windward side, where plant height was lowest (9.4 cm). Grasses were not correlated either positively or negatively with either shrubs or forbs. Percent cover by grasses was not associated with plant height Table 5. Plant mineral concentrations downslope on windswept communities. Minerals (ppm) Zinc Slope position windward side Slope Top position leewa: Middle rd side Top 14.0 Middle 23.7 Bottom 25.0 Bottom 15.7 20.7 23.7 Iron 308. 491. 627. 379. 366. 706. Phosphorus 1,400. 1,900. 2,200. 1,500. 1,900. 2,400. Manganese 35. 38. 70. 45. 67. 40. Copper Sodium 10. 226. 9. 250. 11. 287. 8. 237. 9. 249. 11. 287. Nitrogen Calcium 20,000. 21,000. 21,000. 20,000. 23,000. 21,000. 7,800. 9,400. 15,400. 9,300. 10,000. 8,700. Magnesium Potassium 1,800. 13,700. 2,400. 13,600. 2,800. 14,500. 2,100. 13,.300. 2,300. 13,400. 2,000. 14,200. 88 Great Basin Naturalist Vol. 42, No. 1 as were shrubs and forbs. Forbs as a life form were not positively correlated with any fac- tor and grasses were not negatively corre- lated with any factor. Plant height was neg- atively correlated with density because the highest density was at the top of the ridges where height was lowest. Soil minerals tended to be more positively than negatively associated with the other fac- tors (Table 7B). Grasses were positively cor- related with phosphoms and nitrogen down- slope. Two studies have shown that an increase in phosphorus and nitrogen increases the amount of biomass produced on range- lands (Barrett 1979, Wight 1976). Magnesium was the only soil mineral positively corre- lated with shrubs. Copper was highly corre- lated with several factors in the soil and plants (Table 7B). It was the only mineral to be correlated either positively or negatively between the plant and soil. As above, mineral concentrations in plants were also more positively than negatively correlated with the other factors. Zinc, iron, and phosphorus were positively associated with shrub cover. These minerals increased downslope as did percent cover by shrubs. Forbs showed the reverse trend, with percent cover by forbs decreasing downslope as zinc, iron, and phosphorus increased. Nitrogen in plants was positively correlated with plant height. Vegetative and soil differences in areas subjected to high winds develop character- istics that distinguish them from the sur- rounding plant community. The tops of the windswept ridges have several unique fea- tures (i.e., percent exposed rock, soil depth, plant height, plant cover, and composition of life form). Management of these areas for livestock or wildlife should include special considerations. The windswept ridges during winter are often snow-free while the sur- rounding areas are covered. This would tend to concentrate animals in these areas and cause overgrazing. Also, these areas are the first to green up in the spring due to the ex- posure, which may be why animals graze these areas while plant carbohydrates are low. Improvements or rehabilitation of these areas (i.e., brush control, range reseeding, etc.) by range managers must be looked at closely. Anderson et al. (1976) showed in Wyoming that windswept areas had a differ- ent relative cover by palatability class than the surrounding vegetation type, even though both areas rated fair for range condition. Plant types and species used to improve these areas must be able to withstand the harsh en- vironment of the windswept ridges. Life forms of the plants should show adaptation (i.e., low, mat forming, rhizomatous, drought- and cold-hardy grasses or forbs) able to with- stand the unique environments of the sites. Revegetation attempts in these areas would best be achieved by planting mixtures of seeds rather than using single seed species in reseeding projects. Mixtures would allow greater variability in the plant resource in meeting the needs of an ever-changing and varied habitat. Jaynes and Harper (1978) stated that species of undesirable forage val- ue may be the only plant species available to meet the criteria of areas with harsh environ- ments (such as windswept ridges). Vegetative cover to prevent erosion may have priority over palatable forage in certain areas. Table 6. Plant and soil ratios of mineral concentrations with regard to slope position. Slope position windward side Minerals Slope position leeward side Zinc Iron Phosphorus Manganese Copper Sodium Nitrogen Copper Magnesium Potassium Top Middle Bottom Top Middle Bottom 10.8 16.9 17.9 7.5 12.9 19.8 18.8 45.1 43.2 17.9 17.3 41.5 117.6 175.9 139.2 107,9 137.7 119.4 2.9 3.4 3.9 3.5 4.5 2.1 12.5 10.0 9.2 11.4 15.0 12.2 5.4 5.9 7.0 5.9 7.0 7.0 10.0 15.0 11.7 9.5 11.5 11.1 2.0 1.6 4.1 2.8 3.5 3.4 10.7 4.0 5.7 9.4 10.0 5.9 81.0 58.6 48.3 63.9 67.0 49.3 March 1982 MoRETTi, Brotherson: Vegetation-Soil Factors 89 Management of windswept communities and ranges with roUing hill country, as discussed in this paper, should be preceded by a careful study of the vegetative and soil differences to insure successful management programs. Literature Cited Anderson, D. L., W. K. Ostler, C. Freeman, and K. T. Harper. 1976. Vegetation of the proposed China Butte coal strip mine. Rocky Mountain Energy Co. Report 1976: 4-2.3. Baker, D. E. 1974. Copper: soil, water, plant relation- ships. Federation Proc. ,33:118-119.3. Barrett, M. W. 1979. Evaluation of fertilizer on prong- horn winter range in Alberta. J. Range Manage. 32(l):55-59. Bloss, D., and J. Brotherson. 1979. Vegetation re- sponse to a moisture gradient on an ephemeral stream in central Arizona. Great Basin Nat. 39:161-176. BoYuoucos, G. J. 1951. A recalibration of the hydro- meter method for making mechanical analysis of soils. J. Agron. 43:4.34-4.38. Brotherson, J. D. 1979. Ecological and community relationships of Eriogonum corymbosum (Poly- gonaceae) in the Uinta Basin, Utah. Great Basin Nat. 39:177-191. Brotherson, J. D., and S. T. Osayande. 1980. Mineral concentrations in true mountain mahogany and Table 7. Correlation coefficients of environmental and biotic factors with respect to each other: Section A, life form, general vegetative and site factors; Section B, mineral concentration in the soil; Section C, mineral concentra- tion ill the plants; subscript "p," mineral concentrations in plant; subscript ' .-...^-.- perscripts "2,3,4," significant levels as follows: 2 = 0.05, 3 = 0.02, 4 = 0.01. mineral concentrations in soil; su- Factor A Shrub Forb Grass Annual Plant height Plant density Organic matter pH Soluble salts Soil depth Positive correlations Negative correlations Soil depth^, height* Soil depth^, Zn^p, shrub^ Cu^s, Mg^s, M^ soluble salts*, Zn^p Ca4„ Ca2p, M^p Cu^, Ca^^, Mg*s organic matter^, Zn^p shrub^, height^ Forb^ Zn^p, shrub'* Plant density^ Soil depth^, heigh t^ Fe4„ F^„ Cu3„ Na3„ K^,, Fe^p, P^p, plant Plant density* B Zinc Iron Phosphorus Manganese Copper Sodium Nitrogen Calcium Magnesium Potassium Fe2,, Mn^s Na2 P*s, K^, Na2p Na'*;, Mg*s, K'*s, organic matter^ soluble salts', Zn** Fe^p, Cu^p, Fe^s pH^, soluble salts2, Mg^p Cu'*s, organic matter^, soluble salts4, Zn4 Fe2 F3„ Cu^^, Fe<„ Na^p P^P Ca2s, pH4, M^p, Plant densit)^ Ca2„ ¥?„ pH2 pFP pH3 Fe2_o pH2 C Zinc Iron Phosphonis Copper Sodium Calcium Magnesium Potassium Cu'*s, Mg*5, organic matter*, soluble salts^, P*p, Cu^p, Height^ Cu4„ Na3„ M^„ K^, P*^, Cu2p Cu4„ Na^s, Mg^s. K^p Cu2,, Na2,, Zn-*p, I^,, Mn2,, K2, Mn^s, Mg*s Ca3„ pPP, Mn3p P2„ Zn^p, Fe^p, Cu^p, Fe2p, P*p Ca-* Forb^ pH-* pH3 pH-* Plant density^ Fe2. 90 Great Basin Naturalist Vol. 42, No. 1 Utah juniper and in associated soils. J. Range Manage. .33:182-185. Charley, J. L. 1977. Mineral cycling in rangeland eco- systems. In R. E. Sosebee, ed., Range plant phys- iology. Society for Range Management, Range Science Series No. 4. Daubenmire, R. 1959. A canopy-coverage method of vegetational and analysis. Northwest Sci. 33:43-46. Epstein, E. 1972. Mineral nutrition of plants: principles and perspectives. John Wiley and Sons, New York. 412 pp. Fairchild, J. AND J. D. Brotherson. 1980. Microhabitat relationships of six major shrubs in Navajo Na- tional Monument, Arizona. J. Range Manage a3: 150-156. Greenwood, L. C, and J. D. Brotherson. 1978. Some ecological relationships between pinyon-juniper and birchleaf mountain mahogany stands. J. Range Manage. 31(3): 164-168. Harner, R. F., and K. T. Harper. 1973. Mineral compo- sition of grassland species of the eastern Great Basin in relation to stand productivity. Can. T. Bot. 51:2037-2046. Hess, P. R. 1971. Textbook of soil chemical analysis. Chem. Publishing Co., ew York. Isaac, R. A., and J. D. Kerber. 1971. Atomic absorption and flame photometry: techniques and uses in soil, plant, and water analysis. Pages 17-.38 in L. M. Walsh, ed.. Instrumental methods for analysis of soils and plant tissue. Soil Sci. Soc. Amer. Wis. Jackson, M. L. 1958. Soil chemical analysis. Prentice- Hall, Englewood Cliffs, New Jersey. Jaynes, R. a., and K. T. Harper. 1978. Patterns of natu- ral revegetation in arid southeastern Utah. J. Range Manage. 31:407-411. Jones, J. B. 1973. Soil testing in the United States. Comm. Soil Sci. Plant Anal. 4:307-.322. Levins, R. 1966. The strategy of model building in ecol- ogy. Am. Sci. 54:412-431. Lindsay, W. L., and W. A. Norvall. 1969. Devel- opment of a DTPA micronutrient soil test, Agron. Abstracts. Equilibrium relationships of Zn-t- +, Fe+ -I-, Cu-»- +, and H-l- with EDTA and DTPA in soil. Soil Sci. Soc, Amer. Proc. 33:62-68. MacArthur, R. H. 1972. Geographical ecology: patterns in the distribution of species. Harper and Row, New York. 251 pp. Marinos, N. G. 1978. Fellfields and cushion plants of the Rockies. Utah Sci. 6:68-72. National Range Council. 1976. Nutrient requirements of beef cattle. 5th ed. National Academy of Sci., Washington, D.C. 56 pp. 1976. Nutrient requirements of .sheep. 5th ed. National Academy of Sci., Washington, D.C. 72 pp. Russell, D. A. 1948. A laboratory manual for fertility students, .3d ed. Wm Brown Co., Dubuque, Iowa. 56 pp. Shannon, C. E., and W. Weiner. 1949. The mathemati- cal theory of communication. University of Il- linois Press, Urbana. 65 pp. Sneath, p. H. a., and R. R. Sokal. 1973. Numerical tax- onomy, the principles and practices of numerical classification. W. H. Freeman and Co., San Fran- cisco, California. 547 pp. Wight, J. R. 1976. Range fertilization in the northern Great Plains. J. Range Manage. 29:108-185. WEATHER CONDITIONS IN EARLY SUMMER AND THEIR EFFECTS ON SEPTEMBER BLUE GROUSE {DENDRAGAPUS OBSCURUS) HARVEST Joy D. Cedarleaf , S. Dick Worthen', and Jack D. Brotherson' Abstract.- Relationships of temperature and precipitation to the reproductive success of blue grouse {Dendra- gapus obscunts) were investigated. Maximum and minimum temperatures followed similar patterns during the years 1976-1981 and showed no patterns relative to hatching success. Precipitation data, however, was variable. When sig- nificant amounts of precipitation fell during the last three weeks of the hatching period, chick survival and, there- fore, recruitment were adversely affected. We suggest that precipitation occurring at the end of the hatch period re- duces the September harvest of birds. Blue grouse {Dendragapus obscurus) in- habit fir forests in many western states (Beer 1943). The range of this species is closely as- sociated with the distribution of true fir (Abies) and Douglas fir (Pseudotsuga) in west- em North America (Beer 1943). Blue grouse depend on conifer cover in winter and on shrubs, forbs, and grasses during the spring and summer (Rogers 1968). Most blue grouse are migratory, moving from spring and sum- mer ranges in open meadows to timbered wintering areas at higher elevations in winter (Johnsgard 1973). The quality of breeding habitat seems to affect the number of breeding birds (Zwickel et al. 1968). In the latter study, grazed and imgrazed habitats and their effects on blue grouse breeding success were compared. There was greater breeding success on un- grazed areas (Zwickel et al. 1968). Her- baceous vegetation that suffers under heavy grazing is used by the birds for food and cov- er. Good grazing practices are important for the maintenance of blue grouse populations (Mussehl 1963). Zwickel and Bendell (1967) suggest that another important factor in regulating the population densities of blue grouse is the dis- persal of the juvenile birds to winter ranges. They also noted that temperature and mois- ture did not seem to affect the two broods they observed. As a species, blue grouse have become im- portant upland game birds in the state of Utah in recent years (Bunnell et al. 1977). Af- ter reestablishing and lengthening the hunt- ing season, efforts are presently being made to determine the general status of popu- lations as well as population trends (Rogers 1963). Because of their solitary nature and the dense cover they inhabit, their popu- lation densities are difficult to estimate (Johnsgard 1973). Fluctuating population densities and inconsistencies of hatch makes management and utilization of the blue grouse resource difficult. In this study, tem- perature and precipitation were compared to the hatching success of blue grouse to see if either of these factors affected the age and number of birds harvested. Study Site The study area lies along the west slopes of the Wasatch Mountains of central Utah (the Wasatch Front), extending from American Fork Canyon on the north to Hobble Creek Canyon on the south. This represents a dis- tance of some 50 km (30 m). Here several major canyons and their drainage basins ex- tend eastward into the mountains away from the Wasatch Front. Coniferous forests (sub- alpine fir zone), in which Douglas fir and species of true fir are prominent, occur in the drainage basins and along the canyon slopes. The meadow and open areas associated with this zone contain shrubs such as chokecherry {Primus spp.), serviceberry {Amelanchier 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 'Division of Wildlife Resources-State of Utah, 1115 North Main, Springville, Utah 8466.3. 91 92 Great Basin Naturalist Vol. 42, No. 1 spp.), snowberry (Symphoricarpos spp.) and elderberry {Sambucus spp.), as well as vari- ous forbs and grasses. Materials and Methods Ten weather stations were chosen to repre- sent the study area. These stations were all located in Utah County, Utah, and varied in elevation from 1370 m (4497 ft) to 1720 m (5640 ft). The data collected at these stations were obtained from the National Climatic Center in Asheville, North Carolina. The maximum and minimum temperatures of each week for the months of May, June, and July we K summed and averaged for each of the ten weather stations. The precipitation data for this time period were also summed and averaged. The weekly precipitation and temperature averages were then plotted against the weekly hatching curve for the same year. This was done for the years 1976-1981. The weather stations located in the same general vicinity were grouped to- gether and those figures were then averaged and plotted against the hatch data. The pre- cipitation received at each station was tallied separately, averaged, and plotted on the graph against the hatch curve (Figs. 1-2). Checking stations were set up to gather harvest information. Wings were used to de- termine the sex and age of each bird. Three stations were located in canyons (American Fork, Provo, and Hobble Creek) that con- tained populations of blue grouse. Each sta- tion was operated from 9:00 a.m. until 7:00 p.m. on each of the two days of the opening weekend of the grouse season. One wing was 10 16 22 28 .Prectp 1978 10 16 22 28 Fig. 1. Temperature and precipitation plotted a^ain,st the number of birds in the hatch curve for the years 1976-1978. ^ March 1982 Cedarleaf et al.: Blue Grouse 93 collected from each blue grouse brought to the station, as well as hunter success informa- tion. In addition to the check stations, wing barrels were set up to gather wings during the first two weeks of the hunt. Wings collected at the stations and the wing barrels were analyzed by Division of Wildlife Resources biologists to determine sex and age of the birds (Bunnell et al. 1977). Additional data were gained from the juve- nile wings by back dating to the time of the hatch (Schladweiler 1970). Results and Discussion The maximum and minimum temperatures generally followed the same pattern each year (Figs. 1-2). None of the years or stations contained extreme maximum or minimum temperatures. The years 1976 to 1979 had little or no precipitation during the latter part of the hatch period, but the years 1980 and 1981 had precipitation during the last two weeks of the hatch period (Figs. 1-2). The hatch curve as well as the percentage of juveniles contained in the harvest were de- termined from the wings collected (Schladweiler 1970). A high number of juve- niles (75 percent) in the harvest indicates good or high production and recruitment into the population. In the years 1980 and 1981, 48 percent of the birds harvested were juve- niles. In the years 1976, 1977, 1978, and 1979 the harvest included 78 percent, 63 percent, 66 percent, and 73 percent juveniles, respec- tively. These values are believed to give good indication as to population composition dur- ing these years. If the percentage of the 1979 Ma« 1980 .Piecip _4 0 1980 10 16 22 28 July qn fin ^ S — 50 60 5 n 40 as 5 S 30 E % ?n 10 16 22 28 10 16 22 28 Fig. 2. Temperature and precipitation plotted against the number of birds in the hatch curve for the years 1979-1981. 94 Great Basin Naturalist Vol. 42, No. 1 harvested juvenile birds is low, then the pop- ulation is considered down. The years 1980 and 1981 had a low percentage of juvenile birds when compared to the years 1976 to 1979. If we graph the precipitation during the last three weeks of the hatch period, and the percentage of juveniles in the harvest against time in years, an interesting relation- ship appears (Fig. 3). The two lines become mirror images of one another. Further, if per- centage of juveniles in the harvest and total precipitation during the last three weeks of the hatch period are graphed against each other (Fig. 4), the relationship shown is statis- tically significant (P < 0.05). As the precipi- tation during this period increases, the per- cent of juveniles in the harvest decreases. This suggests that precipitation late in the hatch period affects chick survival and recruitment. Zwickel (1967) suggests that moisture and cold weather do not affect the chicks. He also suggests that the dispersal of the juve- niles to their winter ranges is the major pop- ulation regulating factor. The dispersal of the juveniles would seem to affect each brood separately and therefore should not have an overall effect on population. Factors which could reduce the proportion of juvenile birds in the harvest could be pre- cipitation, temperature, or a combination of both. Our study suggests that precipitation during the hatching season may be respon- sible for few juveniles in the harvest. Extreme amounts of precipitation or extremely low temperatures during a couple of days late in the hatching period could affect the popu- lation drastically. This affect on the 1 1 1978 1979 Years Fig. 3. Percent juveniles in the harvest plotted against the total precipitation (inches) during the last three weeks of the hatching season for the years 1976-1981. 100 50 • y = -0 036x + 2 84 r = -0 78 Sig = 0,05 2 4 6 .8 10 12 1,4 16 17 1,8 Total precipitation (incties) during last 3 weeks of hatching season Fig. 4. Percent juveniles in the harvest plotted against the total precipitation (inches) during the last three weeks of the hatching season. population may not be shown in our data be- cause it would be masked by the averages of temperature and precipitation. Additional studies are needed to determine the specific regulating factors of blue grouse populations. This study seems to indicate that the amount of moisture received during the last part of the hatching period can affect the number of juveniles entering the population. Literature Cited Beer, J. 1943. Food habits of the blue grouse. J. Wildlife Manage. 7:32-44. BoAG, D. A. 1963. Significance of location, year, sex, and age to the autumn diet of blue grouse. J. Wildlife Manage. 27:555-562. Bunnell, D. S., J. S. Rensel, J. F. Kimball, and M. L. Wolfe. 1977. Determination of age and sex of dusky blue grouse. J. Wildlife Manage. 41:662-666. Fowle, C. D. 1960. A study of the blue grouse on Van- couver Island, B.C. Canadian J. Zool. 38:701-713. Johnsgard, p. a. 1973. Dusky blue grouse. Pages 175-192 in Grouse and quail of North America. Univ. of Nebraska Press. MussEHL, T. W. 1963. Blue grouse brood cover selection and land use implications. J. Wildlife Manage 27:579-554. Rogers, G. B, 1963. Blue grouse census and harvest in the United States and Canada. J. Wildlife Man- age. 27:579-585. 1968. The blue grouse in Colorado. Colorado Game, Fish, and Parks Dept., Game Research Di- vision Technical Publication No. 21. SCHLADWEILER, P., T. W. MuSSEHL, AND R. J. GrEENE. 1970. Age determination of juvenile blue grouse by primary development. J. Wildlife Manage. 34:649-652. Smith, N. D., and I. O. Buss. 1963. Age determination and plumage observations of blue grouse. J. Wildlife Manage. 27:566-578. March 1982 Cedarleaf et al.: Blue Grouse 95 Tervort, p. 1976-1979. Upland game inventory report. Utah Division of Wildlife Re.source.s. P.R. Project W-65-R. WoRTHEN, S. D. 1980 and 1981. Upland game inventory report. Utah Division of Wildlife Resources. P.R. Project W-65-R. ZwiCKEL, F. 1972. Some effects of grazing on blue grouse during summer. J. Wildlife Manage. 36:631-634. 1967. Some observations of weather and brood behavior in blue grouse. J. Wildlife Manage. 31:563-567. ZwicKEL, F., AND J. F. Bendell. 1967. Early mortality and the regulation of numbers in blue grouse. Ca- nadian J. Zool. 45:817-851. Zwickel, F., I. O. Buss, and J. H. Brigham. 1968. Au- tumn movements of blue grouse and their rele- vance to populations and management. J. Wild- life Manage. 32:456-467. DESCRIPTION OF THE FEMALE OF PHALACROPSYLLA HAMATA (SIPHONAPTERA:HYSTRICHOPSYLLIDAE) R. B. Eads' and G. O. Maupin' Abstract.- The previously unknown female of Phalacropsylla hamata Tipton & Mendez is described and an allo- type is designated. A key is provided to aid in distinguishing females of this genus. The genus Phalacropsylla includes an inter- esting assemblage of species adapted to high altitudes. Described species include P. para- disea Rothschild 1915, P. alios Wagner 1936, P. nivalis Barrera & Traub 1967, P. hamata Tipton & Mendez 1968, and P. oregonensis Lewis & Maser 1978. With the exception of alios, these nest fleas have not been com- monly collected, even during extensive ecto- parasite studies (including nest examinations) within the known range of the genus in the western United States and Mexico. Host re- cords suggest that wood rats, Neotonia spp., and closely associated rodents and la- gomorphs are normal hosts of the Phalacropsylla. Phalacropsylla hamata was described from a single male collected by V. J. Tipton from a rodent nest in the Sierra Madre Occidental Range (Cerro Potosi) Nuevo Leon, Mexico, 20 April 1964, at an elevation of 3050 m. We have recently received several specimens referable to this species from the vicinity of Albuquerque, New Mexico. Phalacropsylla is redescribed to summarize the characteristics of this genus. Minor differences between the New Mexico males and the male holotype of hamata probably represent individual varia- tion rather than subspecific differences (Fig. 3-5). Phalacropsylla Rothschild 1915 Frontal tubercle and striarium absent. Eyes vestigial, eye spot lightly sclerotized. Genal comb of 2 overlapping teeth; outer tooth 'Vector-Borne Viral Diseases Division. Center for Infectious Diseases, Centers for Disease Control, Public Health Service, U.S. Department of Health and Human Services, P.O. Box 2087, Fort Collins, Colorado 80522-2087. slightly over V2 length of narrower, subacute inner tooth. Pronotal comb of 14-18 spines. Abdominal spinelets and mesonotal pseudo- setae present. Some subapical bristles of in- ner side of hind coxae weakly spiniform. Fifth segment of all tarsi with 4 lateral pair of bristles and 1 pair shifted to the plantar surface between the 1st lateral pair. Sternum of male V-shaped, distal arms bifid; distal arm with lightly sclerotized dorsal expansion and short, preapical spiniform bristles, plus long, curved, spiniform bristles in certain species. Phalacropsylla hamata Tipton & Mendez Phalacropsylla hamata Tipton & Mendez, 1968, Pac. In- sects 10: 177-214. Material.— All from Bernalillo Co., New Mexico, collected by Curt Montman. Allo- type female ex Peromyscus leucopus, 20 No- vember 1980. Parallotype female with same data. Male collections include 2 ex P. leuc- opus 20 November 1980, 1 ex Neotoma albi- gula 20 November 1980, and 1 ex N. albigula 20 February 1981. Allotype deposited in the United States National Museum of Natural History collection, Washington, D.C. Diagnosis.— Flea specific differentiation in the absence of males is often difficult or impossible. In the Phalacropsylla, however, configuration of the caudal lobe of sternum VII seems to show sufficient distinctiveness to be of value in female specific identi- fication. In hamata, paradisea, and orego- nensis the female sternum caudal lobe is 96 March 1982 Eads, Maupin: Siphonaptera 97 broader than long. The lobe is ca 3.5 X as broad as long in hamata, as opposed to less than 2 X as broad as long in paradisea and oregonensis. PhalacropsyUa males are readily identified to species with the key presented by Lewis and Maser (1978). Description of female.— Head: Pre- antennal region with 2 fairly straight rows of bristles. Frontal row of 4 small, thin bristles; ocular row of 4 much larger ones and a fine bristle slightly out of line and cephalad of eye spot; 3 thin bristles caudad of ocular row. Maxilla narrow, acuminate distally, extending to base of 4th segment of maxillary palpus; maxillary palpus extending ca % length of coxa I. Labial palpus extending beyond apex of coxa L Postantennal region with bristles arranged 1:3:5 on one side and 1:4:5 on the other, the caudal row with fine intercalaries; 18-20 fine hairs along the antennal fossa. TJiorax: Pronotum with a row of 6 large bristles, ventrad bristle 2 X length of others, separated by 5 smaller bristles per side; pro- notal com b of 14 spines of ca same length, except for shorter ventralmost pair. Mesono- tum with a row of 5 large bristles and fine in- tercalaries preceded by a row of 7 smaller ones and 15-18 fine bristles scattered along cephalad margin; mesonotal flange with 2 pseudosetae per side (parallotype with 3). Mesepisternum with a large, lateral bristle (parallotype with a fine bristle preceding large bristle on 1 side); mesepimeron with 2 irregular rows of bristles arranged 3:2 on one side and 4:2 on the other. Metanotum (exclu- sive of lateral metanotal area) with 3 rows of bristles, caudal row of 4-5 large ones and 5 fine intercalaries, a median row of 6 or 7 smaller bristles and a cephlad row of 2 or 3 still smaller bristles; a single fine bristle pre- cedes the 3 rows; lateral metanotal area with 1 large dorsocaudal bristle, a large caudal bristle, and a fine ventral bristle. Mete- pisternum with 3 subdorsal bristles in a row, a large one flanked by 2 smaller ones; mete- pimeron with ca 6 lateral bristles arranged 3:3:1 (paratype 2:3:1). Abdomen: Terga I-IV with apical spinlets arranged (3-4), (4-4), (2-1), and (1-0), paral- lotype (3-3), (4-3), (3-3), and (1-1). Terga typi- cally with median row of 5-7 large bristles alternating with smaller ones extending al- most to ventral margin, preceded by a some- what shorter row of 7 or 8 smaller bristles and 3 or 4 shorter bristles in an irregular row or patch on dorsocephalad margin. Middle antepygidial bristle ca 2 X length of ventral bristle and almost 3 X length of dorsal bristle. Sternum II with a vertical row of 3 large bristles, preceded by irregular row of 5 I '^ 3 Figs. 1-3. Phalacropsylla hamata. 1, female sternum VII; 2, spermatheca; 3, male sterna VIII, IX. 98 Great Basin Naturalist Vol. 42, No. 1 penis rods crochet sclerotized inner tube fulcrol latero- ventrol lobe immovable process movable process Figs. 4,5. Phalacropsylla hamata. 4, male aedeagus; 5, male clasper. March 1982 Eads, Maupin: Siphonaptera 99 or 6 smaller bristles and 3 or 4 small bristles toward ventral margin; other unmodified sterna with 4 large bristles preceded by 3 or 4 smaller ones. Modified abdominal segments: (Fig. 1,2). Sternum VII with a large lobe on ventral half or caudal margin ca 3.5 X as broad as long; sinus directed cephalad, not forming definite ventral lobe. Sternum VII with subventral row of 4 (or 5) large bristles in a slightly oblique row, preceded by an irregular row of 7 or 8 smaller bristles. Spermatheca with bul- ga ca 1.5 X as broad as high; dorsal and ven- tral margins slightly concave at basal 3rd, be- coming convex to form dilated, caudal % of bulga; hilla upcm-ved at basal 3rd, over 4 X as long as wide, about same width through- out, no constriction at basal 3rd. Tergum VIII with 2 irregular rows or 7 of 8 small, thin, bristles above the 8th spiracle; a curved row of 8-10 large bristles nearly sinuate, caudal margin, row beginning beneath ven- tral anal lobe and terminating just above ven- tral margin; 3-4 smaller, marginal bristles be- low ventral anal lobe, with 5-7 small. scattered, submedian bristles anterior to above fringe row. Discussion The New Mexico P. hamata collection site was the rocky, lower slopes (1600-1800 m) of the Sandia Mountains, east of Albuquerque. These fleas were taken at elevations that in- clude grassland communities grading into juniper {Juniperus monosperma)-pinyon {Pinus edulis) woodland. Hosts included Per- omijscus leucopus, which lives among apache plume {Falliigia paradoxa) and saltbush {Atri- plex canescens), and Neotoma alhigula, which is found in both grassland and juniper-pinyon communities (Findley et al. 1975). Acknowledgments We thank Dr. Robert Traub, University of Maryland, for comparing our material with the holotype of P. hamata in the U.S. Nation- al Museum collection. Mr. Curt Montman, Bernalillo County Department of Health, New Mexico, kindly provided the specimens. 1. 2(1). 3(1). 4(3). Key to female PhalacropsijUa Caudal lobe of st VII longer than broad 2 Caudal lobe of st VII broader than long 3 Caudal lobe of St VII ca 1.5 X as long as broad nivalis Caudal lobe of st VII ca 1.9 X as long as broad alios Caudal lobe of st VII 3.5 X as broad as long hamata Caudal lobe of st VII less than 2 X as broad as long 4 Caudal lobe of st VII rectangular, broadly rounded at apex paradisea Caudal lobe of st VII more triangular, apex bluntly pointed and deflected ven- trally oregonensis Literature Cited Findley, J. S., A. H. Harris, D. E. Wilson, and C. Jones. 1975. Mammals of New Mexico. Univ. of New Mexico Press, Albuquerque, New Mexico. 260 pp. Lewis, R. E., and C. Maser. 1978. PhalacropxiiUa orego- nensis sp. n., with a key to the species of Phala- cropsijUa Rothschild 1915 (Siphonaptera: Hys- trichopsyllidae). J. Parasit. 64(1): 147-150. Tipton, V. J., and E. Mendez. 1968. New species of fleas (Siphonaptera) from Cerro Potosi, Mexico, with notes on ecology and host parasite relation- ships. Pac. Insects l6(l): 177-214. FIRST RECORD OF PYGMY RABBITS {BRACHYLAGUS IDAHOENSIS) IN WYOMING Thomas M. Campbell IIP, Tim W. Clark\ and Craig R, Groves' Abstract.- Pygmy rabbits are reported for the first time in southwestern Wyoming. The range for this species is thus extended 240 km and 145 km from the nearest records in Idaho and Utah, respectfvely. ^ Although the pygmy rabbit occupies much of the sagebrush {Artemisia tridentata) habi- tat of the Great Basin, it has not been pre- viously reported from Wyoming other than a single, unconfirmed observation described by Green and Flinders (1980a). Between 11-20 October 1981, 6 specimens were collected (2 males, 4 females), 17 indi- viduals were observed, and 2 skulls and nu- merous pellets were found at two sites in southwestern Wyoming. The first site was in South Fork Muddy Creek drainage, 8 km north and 8 km east of Carter, Uinta County Wyoming (T17N R115W 84,5,6,33- TUN R116W Sl,23, 24). The second site was along North Fork Muddy Creek drainage in Cum- berland Flats, Lincoln and Uinta counties, Wyoming (T16N R118W S34; T19N R115W S19; T19N R116W S2,14). The two sites were geographically separated by the Hogs- back and Oyster Ridge highlands. The nearest previously reported records of pygmy rabbits are from the west near Poca- tello, Bannock County, Idaho (Davis 1939), and near Clarkston, Cache County, Utah (Durrant 1952). These two new locales ex- tend the known range of B. idahoensis ap- proximately 240 km to the southeast and 145 km to the northeast, respectively. In view of the rough mountainous topography and the apparent lack of habitat between known range and this range extension, the pygmy rabbits in Wyoming may be a disjunct population. Our observations at both sites showed B. idalioensis primarily confined to dense stands of big sagebrush growing in deep soils of drainages and hollows. This concurs with findings reported by Grinnell et al. (1930), Fisher (1979), and Green and Flinders (1980b). We also observed sign and collected two animals in a mixed sagebrush- greasewood {Sarcobatus spp.) habitat similar to that described by Davis (1939). Mean external measurements of collected specimens were as follows: Females (n = 4), total length, 265 mm; tail length, 23 mm; hind foot length, 78 mm; ear length, 53 mm. Males (n = 2), total length, 238 mm; tail length, 21 mm; hind foot length, 77 mm; ear length, 55 mm. The authors are presently engaged in a de- tailed study of distribution, habitat relation- ships, and taxonomic status of pygmy rabbits in Wyoming. We acknowledge William Oelklaus for suggesting possible occurrence of this species in Cumberland Flats. Field assistance in an earlier unsuccessful attempt to document pygmy rabbit presence in Cumberland Flats was provided by Mark Stromberg and How- ard Hunt. Denise Casey provided field assist- ance and critical review of the manuscript. Literature Cited Davis, W. B. 1939. The recent mammals of Idaho. Cax- ton Printers, Caldwell, Idaho. 400 pp. Durrant, S. D. 1952. Mammals of Utah. Univ. of Kansas Publ. Mus. Nat. Hist. 6. .549 pp. Fisher, J. W. 1979. Reproduction in the pygmy rabbit in southeastern Idaho. Unpublished thesis. Idaho State Univ., Pocatello. 32 pp. Green, J. S., and J. T. Flinders. 1980a. Brachylagus idahoensis. Mammal. Species No. 125. Amer. Soc. of Mammal. 4 pp. 1980b. Habitat and dietary relationships of the pygmy rabbit. J. Range Manage. .33:136-142. Grinnell, J., J. Dixon, and J. M. Linsdale. 1930. Ver- tebrate natural history of a section of northern California through the Lassen Peak region. Univ. of California Pub. Zool. .35. 594 pp. 'Biota Research and Consulting, Inc., Box 2705, Jackson, Wyoming a300L 'Department of Biology, Idaho State University, Pocatello, Idaho 83209. 100 PASPALUM DISTICHUM L. VAR. INDUTUM SHINNERS (POACEAE)' Kellv Wayne Allred- .\bstract.- Glabrous Paspalum distichiim, pubescent var. indutum, and pubescent plants of P. distichum from the western United States were compared. No morphological differences other than pubescence exist between the three forms. Plants from the eastern United States are generally more glabrous than those from western regions. The var. indittum represents an extreme pubescent form and is reduced to synonymy under P. distichum. Paspalum distichum L.' is the familiar "knotgrass" of swamps, swales, marshes, ditches, and muddy sites throughout much of the continental United States. It grows along both eastern and western coasts and through- 1 Node Pubescence 2 Collar Hair Length out the southern states. Knotgrass is also widespread in other warm-temperate to trop- ical regions of the world. Material from Dallas County, Texas, with strongly hirsute sheaths and blades was 9 Abaxial Blade Hair Density 10 Panicle Branch Number 3 Sheath Margin Hair Density 4 Sheath Back Hair Density 11 Panicle Internode Length 12 Branch Length 5 Ligule Length 6 Blade Length 13 Rachis Width 14 Spikelet Length 7 Blade Width Adaxial Blade Hair Density 15 Spikelet Width 16 Glume 1 Length Figs. 1-8. Comparisons of the means (vertical line), range (horizontal line), and one standard deviation on each side of the mean (horizontal bar) for three forms of Paspalwn distichium. D, glabrous distichum form; I, pubescent indutum form; W, pubescent western form. The features measured are as labeled. Figs. 9-16. Comparisons of the means (vertical line), range (horizontal line), and one standard deviation on each side of the mean (horizontal bar) for three forms of Paspalum distichum. D, glabrous distichum form; I, pub- escent indutum form; W, pubescent western form. The features measured are as labeled. 'Journal Article 90.3, Agricultural Experiment Station. New Mexico State University, Las Cruces, New Mexico 88003. ^Department of .'Vmmai and Range Sciences, New Mexico State University, Las Cnices, New Mexico 88003. There is current controversv concerning the application of this name, involving also the names P. vaginatum Sw. and P. pa^alodes (Michx.) Scribner. Until this matter is resolved, I will use -p distichum" in the sense of Hitchcock (1951). See also Guedes (1976) and Renvoize and Clayton (1980). 101 102 Great Basin Naturalist Vol. 42, No. 1 described by Shinners (1954) as Paspalum dis- Hchum var. indutum. The variety indutum has been known only from the type locality and a few collections in the same county. Many plants of P. distichum from the western United States, however, including the inter- mountain region, also possess varying degrees of pubescence on the sheaths and elsewhere. This fact suggested the following queries. How does the var. indutum compare with the typical glabrous material of var. dis- tichum, and with the pubescent material of the western United States? Are there other features in addition to pubescence that serve to distinguish any of the three forms? What is the geographic distribution of these forms? How many taxa are involved in this complex? Materials and Methods Plant specimens were gathered from vari- ous herbaria throughout the United States range of Paspalum distichum to give a repre- sentative sample of this taxon. From this sample, 235 specimens were examined and measurements taken for the following fea- tures: node pubescence, collar pubescence (length and number of hairs), sheath margin pubescence (density), sheath back pubescence (density), ligule length, blade length, blade width, adaxial blade pubescence (density), abaxial blade pubescence (density), number of inflorescence branches, length of the up- permost internode of the inflorescence, up- permost branch length, rachis width, spikelet length, spikelet width, and first glume length. Pubescence "density" was measured by the number of hairs touching or intersecting a standard 1 cm micrometer grid at 40X mag- nification. To help evaluafe the relative de- gree of pubescence, a Pubescence Index (PI) was calculated for each specimen. The PI was the sum of the values for the pubescence features (node, collar, sheath margin, sheath back, adaxial blade, and abaxial blade). Total- ly glabrous plants would have a PI of zero. 18.0 10.4 Fig. 17. Pubescence Index values of selected species of Paspalum distichum based on measurements of node pu- axlTbS.'h ]'"''' '"?' f ri? ""f ^'" ^^'' ^'"^"'y' '^'""'^ ^^'^ ^^'' ^^"^i^y- ^d^^'^' blade hair density, and ab- axial blade hair density. Circled PI values represent the indutum form. The map is not intended to represent the ge- ographic distribution of P. disfic/j urn. ^ cm u.c ge March 1982 Allred: Paspalum (Poaceae) 103 with greater PI vakies corresponding to in- creased pubescence. Results and DiscliSsion Most characteristics of Paspalum distichiirn are remarkably consistent throughout the continental United States. The three forms (glabrous distichum, indittum, and pubescent western) are essentially identical for all mor- phological features other than pubescence (Figs. 1-16). Even slight differences, such as the length of the panicle in ternode (Fig. 11) are not correlated with any other features. The obvious differences illustrated by Figures 1-16 are in the pubescence features, but these are the result of the a priori classifica- tion of the plants into three groups: glabrous distichum, pubescent indutum, and pu- bescent western forms. The only possible differences, then, involve pubescence, which can be evaluated by the Pubescence Index (PI). The PI values for 161 specimens of knotgrass were plotted geo- graphically (Fig. 17). (The remaining 74 specimens of the original 235 represented ei- ther duplicates or other specimens with iden- tical PI values and from the same localities as those plotted on the map.) Specimens that were referable to the variety indutum, that is, with densely hirsute sheaths and blades as in the type specimen {Shinners 10564, SMU), were found in widely separate localities from Texas to Washington and had a PI range of 25-79. The type specimen of indutum had a PI value of 74. There appears to be a rough cline in pu- bescence from more glabrous eastern plants (characteristic of distichum) to more pu- bescent western forms. Division of the United States into regions showed an average in- crease in PI values from 2.8 for eastern knot- grass to 18.0 for western plants (Fig. 17). The extreme pubescent forms representing varie- ty indutum showed no such interrelationship with geographic distribution. High PI values Fis 18 Pubescence Index values of selected species of Paspalum distichum based on measurements of sheath back hair density, adaxial blade hair density, and abaxial blade hair density. Circled PI values represent the mdutum form. The map is not intended to represent the geographic distribution of P. distichum. 104 Great Basin Naturalist Vol. 42, No. 1 (7-8) in eastern plants were due mainly to very pubescent nodes, collars, and sheath margins, but the pubescence of western plants involved not only the nodes, collars, and sheath margins, but also especially the sheath back and leaf blades. This is illustrated in Figure 18, where modified PI values from only the sheath back and blade surfaces are plotted. We see that eastern plants are glabrous or nearly so for these features (PI = 0.5), but western plants are often pu- bescent (PI = 9.5). The relationship, however, is far from absolute, as is shown by the range in PI for each region. In conclusion, there is no morphological distinction between the nearly glabrous east- em knotgrass, the pubescent western forms, and the variety indutum. The variety in- diitum merely represents the extreme pu- bescent form of Paspalum distichum. Recog- nition of this taxon is untenable, with no basis in geographic distribution and no correlation with any other suite of features. The limits of such a taxon would be entirely arbitrary. Pubescence patterns in knotgrass do appear in an east-west continuum, but there is no strong geographic distinction nor any feature other than pubescence that would allow a taxonomic segregation. The variety indutum and other pubescent plants are best viewed as pubescent forms of Paspalum distichum. Acknowledgments Many thanks to the curators of the follow- ing herbaria for their generous loan of speci- mens: BRY, FSU, NCU, SMU, UNM, UTC, and WTU. Literature Cited GuEDES, M. 1976. The ease of Paspalum distichum and against futile name changes. Taxon 25(4):512. Hitchcock, A. S. 1951. Manual of the grasses of the United States. USDA Misc. Publ. 200. 2d rev. ed. Agnes Chase. Renvoize, S. a., and W. D. Clayton. 1980. Proposal to reject the name Paspalum distichum L. Taxon 29(3/4):339-340. Shinners, L. 1954. Notes on north Texas grasses. Rho- dora 56:31-32. LOCAL FLORAS OF THE SOUTHWEST, 1920-1980: AN ANNOTATED BIBLIOGRAPHY Janice E. Bowers' Abstract.- Local floras, that is, plant lists for relatively small areas, are widely scattered, often unpublished, and difficult to locate. Over 100 local floras from the southwestern United States (Arizona, Colorado, Nevada, New Mexi- co, and Utah) are listed and briefly annotated. Much of the southwestern United States is well known botanically. Comprehensive, statewide floristic manuals have been pre- pared for Arizona (Tidestrom and Kittell 1941, Kearney and Peebles 1942, 1960), Col- orado (Harrington 1954), New Mexico (Mar- tin and Hutchins 1981), the Sonoran Desert (Shreve and Wiggins 1964), the Mojave Desert (Jaeger 1941), Utah (Welsh and Moore 1973), and the Intermountain Region (Holm- gren and Reveal 1966, Tidestrom 1925, Cronquist et al. 1972, 1977). Floristic studies on a smaller scale have also contributed to knowledge about plant distribution in the Southwest, but these briefer works are often impublished and difficult to locate. Botanical bibliographies such as those compiled by Ewan (1936) and Schmutz (1978) for Arizona, Hoffman and Tomlinson (1966) for Colorado, Christensen (1967a, 1967b) for Utah, and U.S. Fish and Wildlife Service (1977a, 1977b, 1977c) for Colorado, Utah, and New Mexico have not emphasized local floras. The objective of this paper is to provide a List of readily accessible local floras for the Southwest, including Arizona, New Mexico, Utah, Nevada, and Colorado. Only plant lists intended to be more or less complete floras are included. Lists restricted to single life forms (such as cacti, woody plants, fungi, or ephemerals) are not included, nor are man- uals that cover state-sized regions. Only the most recent flora for an area is listed. Both published and unpublished floras are includ- ed, with the reservation that unpublished lists must be readily accessible, either from the re- sponsible agency or through the interlibrary loan programs of major universities. Floras published before 1920 are not included be- cause they are seldom complete and can be difficult to locate. Arizona floras discussed by Bowers (1981) are included here as bibliogra- phic citations only. Arizona AiTCHisoN, S. W. 1978. Oak Creek Canyon and the red rock country of Arizona: a natural history and trail guide. Flagstaff, AZ: Stillwater Canyon Press. AiTCHisoN, S. W., AND M. E. Theroux. 1974. A biotic inventory of Chevelon Canyon, Coconino and Navajo counties, Arizona. Flagstaff, AZ: U.S. For- est Service, Sitgreaves National Forest. Bennett, P. S. 1978. Vascular plants. Grand Canyon Na- tional Park. Grand Canyon, AZ: Grand Canyon Computer Center. Available from Western Ar- chaeological Center, 1415 N. 6th Ave., Tucson, AZ. BoHRER, V. L., AND M. Bergseng. 1963. An annotated catalogue of plants from Window Rock, Arizona. Window Rock, AZ: Navajo Tribal Museum. Bowers, J. E. 1980. Flora of Organ Pipe Cactus Nation- al Monument. J. Arizona-Nevada Acad. Sci. 15:1-11,33-47. Brotherson, J. D., G. Nebeker, M. Skougard, and J. Fairchild. 1978. Plants of Navajo National Mon- ument. Great Basin Nat. 38:19-30. Burgess, R. L. 1965. A checklist of the vascular flora of Tonto National Monument. J. Arizona Acad. Sci. 3:213-223. Butterwick, M., p. Fischer, D. Hillyard, and D. DucoTE. 1980. Annotated checklist of vascular plants collected in the Harcuvar, Vulture and Skull Valley planning units. Phoenix, AZ: U.S. Dept. of the Interior, Bureau of Land Manage- ment. Available from Bureau of Land Manage- ment, Phoenix District Office, 2929 W. Claren- don Ave., Phoenix, AZ. Butterwick, M., D. Hillyard, and B. Parfitt. 1979. Annotated list of taxa of vascular plants collected 'Office of .\rid Lands Studies, University of Arizona. Tucson, Arizona 8.5721. 105 106 Great Basin Naturalist Vol. 42, No. 1 in the Hualapai-Aquarius environmental impact statement area. Phoenix, AZ: U.S. Dept. of the Interior, Bureau of Land Management. Available from Bureau of Land Management, Phoenix Dis- trict Office, 2929 W. Clarendon Ave., Phoenix AZ. Fay, J. M. 1978. Vegetation and flora of the Patagonia- Sonoita Creek Sanctuary, Patagonia, Arizona. Tucson, AZ: Arizona Natural Heritage Program. Available from Arizona Natural Heritage Pro- gram, 30 N. Tucson Blvd., Tucson, AZ. Halse, R. R. 1973. The flora of Canyon de Chelly Na- tional Monument. Tucson, AZ: Univ. of Arizona. Unpublished thesis. Hazen, J. M. 1978. The flora of Bill Williams Mountain. Flagstaff, AZ: Univ. of Northern Arizona. Unpub- lished thesis. Holland, J. S., W. E. Niles, and P. J. Leary. Vascular plants of the Lake Mead National Recreation Area. Las Vegas, NV: Univ. of Nevada, Dept. of Biological Sciences. Lake Mead Tech. Rep. No. 3. Joyce, J. F. 1976. Vegetational analysis of Walnut Can- yon, Arizona. J. Arizona Acad. Sci. 11:127-1.35. Updates Spangle (1953). Keil, D. J. 1973. Vegetation and flora of the White Tank Mountain Regional Park, Maricopa County, Ari- zona. J. Arizona Acad. Sci. 8:35-48. Lane, M. A. 1981. Vegetation and flora of McDowell Mountain Regional Park, Maricopa County, Ari- zona. J. Arizona-Nevada Acad. Sci. 16:29-.38. Lehto, E. 1970. A floristic study of Lake Pleasant Re- gional Park, Maricopa County, Arizona. Tempe, AZ: Arizona State Univ. Unpublished thesis. Leithliter, J. R. 1980. Vegetation and flora of the Chiricahua Wilderness Area. Tempe, AZ: Ari- zona State Univ. Unpublished thesis. Lists .300 taxa for 72.8 km^. Reviews history of the area; de- scribes topography, geology, soils and climate. Compares vegetation of study area to that of nearby mountain ranges. Annotations of the plant list include synonymy, common names, relative abundance, vegetation type, habitat, elevational range, phenology, and collection numbers. Pro- vides small-scale topographic and geologic maps. McCuLLOCH, C. Y., AND C. P. Pase. 1968. Pages 77-88 in Checklist of plants of the Three Bar Wildlife Area. Arizona Game and Fish Dept. Wildlife Re- search in Arizona 1967. Phoenix, AZ: Arizona Game and Fish Dept. McDouGALL, W. B. 1959. Typical seed plants of the ponderosa pine zone. Flagstaff, AZ: Museum of Northern Arizona. Bulletin No. 32. 1959. Plants of the Glen Canyon area in the her- barium of the Museum of Northern Arizona. Flagstaff, AZ: Museum of Northern Arizona. Available from Museum of Northern Arizona Flagstaff, AZ. McDouGALL, W. B. 1962. Seed plants of Wupatki and Sunset Crater National Monuments. Flagstaff, AZ: Museum of Northern Arizona. Bulletin No 37. McDouGALL, W. B., AND H. S. Haskell. 1960. Seed plants of Montezuma Castle National Mon- ument. Flagstaff, AZ: Museum of Northern Ari- zona. Bulletin No. 35. McGill, L. a. 1979. Vascular flora. Pages 56-83 in W. L. Minckley; Sommerfeld, M. R., eds.. Resource inventory for the Gila River Complex, eastern Arizona. Safford, AZ: U.S. Dept. of the Interior, Bureau of Land Management. Available from Bu- reau of Land Management, Safford District Of- fice, 425 E. 4th St., Safford, AZ. McLaughlin, S. P. 1978. Productivity of the understory community in an Arizona ponderosa pine forest. Tucson, AZ: Univ. of Arizona. Unpublished dissertation. McLaughlin. S. P., and W. Van Asdall. 1977. Flora and vegetation of the Rosemont area. Pages 64-98 in R. Davis, and J. R. Calahan, eds.. An en- vironmental inventory of the Rosemont area in southern Arizona. Vol. 1. Tucson, AZ: Univ. of Arizona. Deposited at Univ. of Arizona, Science Library, Tucson, AZ. Pase, C. P., and R. R. Johnson. 1968. Flora and vegeta- tion of the Sierra Ancha Experimental Forest, Arizona. Fort Collins, CO: U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station. Research Paper No. 41. Paulik, L. a. 1979. A vascular flora of the subalpine spruce-fir forest of the San Francisco Peaks, Ari- zona. Flagstaff, AZ: Northern Arizona Univ. Un- published thesis. Petrified Forest National Park. 1976. Seed plant checklist of Petrified Forest National Park. Avail- able from Petrified Forest National Park, Arizona 86028. Phillips, A. M. III. 1975. Flora of the Rampart Cave area, lower Grand Canyon, Arizona. J. Arizona Acad. Sci. 10:148-159. Pierce, A. L. 1979. Vegetation and flora of Buckeye Hills Recreation Area, Maricopa County, Ari- zona. Tempe, AZ: Arizona State Univ. Unpub- lished thesis. Lists 175 taxa for 1500 ha. Reviews history of the area; describes climate, topogra- phy, geology, and soils. Discusses vegetation, list- ing important species for each plant community. Compares flora to that of six other regional parks in Maricopa County. Annotations of the plant list include plant community, relative abundance, and regional distribution. Provides soil and topo- graphic maps, photographs of plant communities. Reeves, T. 1976. Vegetation and flora of Chiricahua Na- tional Monument, Cochise County, Arizona. Tempe, AZ: Arizona State Univ. Unpublished thesis. ScHAAK, C. G. 1970. A flora of the arctic-alpine vascular plants of the San Francisco Mountain, Arizona. Flagstaff, AZ: Northern Arizona Univ. Unpub- lished thesis. Schilling, M. A. 1980. A vegetational survey of the Vol- unteer and Sycamore Canyon region. Flagstaff, AZ: Northern Arizona Univ. Unpublished thesis. Volunteer and Sycamore Canyons are located in southern Coconino County. ScHMUTZ, E. M., C. C. Michaels, and B. I. Judd. 1967. Boysag Point: a relict area on the North Rim of the Grand Canyon in Arizona. J. Range Manage. 20:36.3-369. Simmons, N. M. 1966. Flora of the Cabeza Prieta Game Range. J. Arizona Acad. Sci. 4:93-104. March 1982 Bowers: Southwest Flora Bibliography 107 Smith, E. L. 1974. Phelps Cabin Research Natural Area. Pages 127-13.3 in E. L. Smith, Established natu- ral areas in Arizona; a guidebook for scientists and educators. Phoenix, AZ: Office of Economic Planning and Development. Spangle, P. F." 1953. A revised checklist of the flora of Walnut Canvon National Monument. Plateau 26:86-88. SuNDELL, E. G. 1974. Vegetation and flora of the Sierra Estrella Regional Park, Maricopa County, Ari- zona. Tempe, AZ: Arizona State Univ. Unpub- lished thesis. TooLiN, L. J. 1979. A floral survey of Thomas Canyon, Baboquivari Mountains, Pima County, Arizona. Tucson, AZ; Arizona Natural Heritage Program. Available from Arizona Natural Heritage Pro- gram, 30 N. Tucson Blvd., Tucson, AZ. 1980. Final report on the flora of Ramsey Can- yon. Tucson, AZ: Arizona Natural Heritage Pro- gram. Available from Arizona Natural Heritage Program, 30 N. Tucson Blvd., Tucson, AZ. Ram- sey Canvon is in the Huachuca Mountains, Coch- ise County. TooLiN, L., T. R. Van Devender, and J. M. Kaiser. 1980. The flora of Sycamore Canyon, Pajarito Mountains, Santa Cruz County, Arizona. J. Ari- zona-Nevada Acad. Sci. 14:66-74. Turner, R. M. 1977. Plant species list: Tumamoc Hill. Tucson, AZ; U.S. Geological Survey. Available from USGS Research Project Office, 301 W. Con- gress, Tucson, AZ. Tumamoc Hill is in the Tuc- son Mountains, Pima County. Wadleigh, R. 1969. Plant list for Tucson Mountain Dis- trict of Saguaro National Monument. Tucson, AZ; Saguaro National Monument. Available from Saguaro National Monument, Rt. 9, Box 595, Tucson, AZ. Warren, P. L., and L. S. Anderson. 1980. Annotated checklist of plants of the George Whittell Wild- life Preserve. Pages 80-124 in T. B. Johnson, ed., 1980 progress report for the biological survey of the George Whittell Wildlife Preserve, Graham and Pinal counties, Arizona. Tucson, AZ; Arizona Natural Heritage Program. Available from Ari- zona Natural Heritage Program, 30 N. Tucson Blvd., Tucson, AZ. Welsh, S. L., N. D. Atwood, and J. R. Murdock. Kai- parowits flora. Great Basin Nat. 38:125-179. Yatskievych, G. a. 1980. Plant list for O'Donnell Can- yon, Canelo Hills, Santa Cruz County, Arizona. Tucson, AZ; Arizona Natural Heritage Program. Available from Arizona Natural Heritage Pro- gram, .30 N. Tucson Blvd., Tucson, AZ. Yatskiev\ch, G. a., and C. E. Jenkins. 1981. Fall vege- tation and zonation of Hooker Cienega, Graham Countv, .\rizona. J. .Arizona-Nevada Acad. Sci. 16:7-11. Colorado Barrell, J. 1969. Flora of the Gunnison Basin, Gun- nison, Saguache, and Hinsdale counties, Colo- rado: a study in the distribution of plants. Lists 1062 taxa. Annotated with detailed distributional information. Includes glossary of botanical terms. Fide Donald J. Pinkava, Arizona State Univ. Bradley, R. A. W. 1950. The vascular flora of Moffat County, Colorado. Boulder, CO; Univ. of Colo- rado. Thesis. Lists 511 taxa for 1,231,606 ha; an- notations of the plant list include collector, col- lection location and number, elevation, and habitat. Describes topography, history of plant collection; discusses vegetation zones. Lists plants occurring in Colorado only in Moffat County. Includes location map of study area and photos of vegetation types. Colorado National Monument. 1980. Plants recorded from Colorado National Monument. Fruita, CO: Colorado National Monument. Available from Colorado National Monument, Fruita, CO. Lists 347 taxa for 8276 ha; plant list annotated with common names. Cox, C. F. 1933. Alpine plant succession on James Peak, Colorado. Ecol. Monographs 3;299-.372. Lists 292 taxa. Discusses geology and physiography, cli- mate, effects of grazing and fire, and history of botanical collection. Describes plant commu- nities, with species lists for each. Di.scu.sses en- demic and disjunct species, floristic affinities of the flora, and ecology of alpine plants. Hlina, p. 1980. Preliminary list of vascular plants. Great Sand Dunes National Monument. Alamosa, CO; Great Sand Dunes National Monument. Avail- able from Great Sand Dunes National Mon- ument, Box 60, Alamosa, CO. Lists 252 taxa for 15,764 ha; plant list annotated with common names. Holm, T. H. 1923. The vegetation of the alpine region of the Rocky Mountains in Colorado. Mem. Nat. Acad. Sci. 19; 1-45. -Lists 170 taxa; annotations of plant list include relative abundance, habitat, elevation, and collection localities. Discusses geo- graphical distribution of the species, origins of the flora, vegetation zones. Provides location map. Langenheim, J. H. 1955. Flora of the Crested Butte Quadrangle, Colorado. Madrono 13:64-78. Lists 579 taxa for 25,907 ha; annotations of plant list include collection numbers and species collected outside the study area. Discusses geographical af- finities of the flora and describes vegetation zones. Litzinger, W. 1976. Annotated checklist of vascular plants of Cajon Mesa, Colorado-Utah. Pages 387-4.32 in J. C. Winter, ed., Hovenweep 1975. San Jose, CA: San Jose State Univ. Archaeological Report No. 2. Lists 258 taxa. The study area in- cludes Hovenweep National Monument. Annota- tions include common names, habit, local distri- bution, habitat, phenology, and relative abundance. McNeal, D. W. 1976. Annotated checklist of the alpine vascular plants of Specimen Mountain, Rocky Mountain National Park, Colorado. Southwestern Naturalist 20:423-4.35. Lists 178 taxa; annotations of the plant list include relative abundance, col- lection numbers, habitat, elevational range, and phenologv. Discusses topography, climate; de- scribes alpine habitats. 108 Great Basin Naturalist Vol. 42, No. 1 Nelson, R. A. 1976. Plants of Rocky Mountain National Park. Washington, D.C.: Government Printing Office. Available from Rocky Mountain National Park, Estes Park, CO. Lists 850 taxa; annotations include common names, species descriptions, habitat, local distribution, elevational range, eth- nobotanical notes. Provides keys to species and many photographs of plants and habitats. In- cludes location map and glossary of botanical terms. VoRiES, K. C. 1974. A vegetation inventory and analysis of the Piceance Basin and adjacent drainages. Gunnison, CO: Western State College of Colo- rado. Unpublished thesis. Lists 413 taxa for 416,000 ha; annotations of plant list include local distribution, common names, habit, nativity, rare or endangered status. Discusses physiography, climate, and land use. Quantitative analysis of plan t communities is discussed in detail. Provides small-scale vegetation maps, many photographs of plant communities. Welsh, S. L., and J. A. Erdman. 1964. Annotated checklist of the plants of Mesa Verde, Colorado. Brigham Young Univ. Sci. Bull., Bio. Ser. 4(2):l-32. Lists 411 taxa for 51,813 ha; annota- tions of the plant list include collection location and number, collector, common names, relative abundance, habitat, local distribution, phenology, and plants found in prehistoric ruins. Briefly dis- cusses topography and history of plant collection. Weber, W. A. 1976. Rocky Mountain flora. 5th ed. Boulder, CO: Univ. of Colorado Press. Lists 1600 taxa; annotations include common names, rela- tive abundance, habit, and brief species descrip- tions. Reviews botanical collection; describes plant comnumities; discusses plant geography of the Rocky Mountains. Provides keys to species, glossary of botanical terms; includes small-scale location map and photographs and line drawings of many species. Nevada Beatley, J. C. 1976. Vascular plants of the Nevada Test Site and central-southern Nevada: ecologic and geographic distributions. Springfield, VA: Nation- al Technical Information Center. Lists 1093 taxa for 24,500 km^; annotations include relative abundance, habitat, plant community, common names, local distribution, elevational range, habit, and phenology. Discusses history of bo- tanical collection; describes physiography, geolo- gy, soils, and climate. Describes plant commu- nities, listing dominants. Correlates plant communities with topography and environmental variables. Includes many photographs of plant communities, small-scale maps showing phys- iographic and political features and vegetation types. Bell, K. L., and R. E. Johnson. 1980. Alpine flora of the Wassuk Range, Mineral County, Nevada. Madroiio 27:25-35. Lists 70 taxa for 260 ha; an- notations of plant list include habitat, relative abundance, and regional distribution. Discusses geology, defines alpine zone. Discu.sses endemism and species diversity, compares flora to that of other Great Basin alpine regions. Analyzes geo- graphic affinities of the flora. Bradley, W. G. 1967. A geographical analysis of the flora of Clark County, Nevada. J. Arizona Acad. Sci. 4:151-162. Lists 887 taxa. Discusses geo- graphical affinities and origins of the flora. Clokey, I. 1951. Flora of the Charleston Mountains, Clark County, Nevada. Berkeley, CA: Univ. of California Press. Lists 699 taxa for 170,000 ha; annotations include habitat, associated species, phenology, local distribution, geographical range, elevational range, collector and collection num- ber, synonymy, type locality. Describes botanical collection, geology, climate; discusses endemism, geographical affinities of the flora. Contains keys to species. Freeman, J., and J. L. Mahoney. 1977. Geothermal areas in Nevada: the distribution of vascular plants near the thermal springs surveyed. Ment- zelia 3:8-14. Lists 125 taxa; annotations of plant list include collection locations, relative abun- dance. Includes small-scale location map. Holmgren, A. H. 1942. Handbook of the vascular plants of northeastern Nevada. Logan, UT: Utah State Agricultural College. Lists 1205 taxa; annotations include common names, forage value, local and regional distribution, habitat, relative abundance, synonymy, and elevational range. Discusses bo- tanical exploration, climate, physiography. In- cludes keys to species. Lewis, M. C. 1976. Flora of Santa Rosa Mountains. Og- den, Utah: U.S. Forest Service. Available from Intermountain Region, U.S. Forest Service, Og- den, UT. Lists .347 taxa; annotations of plant list include plant community and collection location. Describes plant communities. Linsdale, M. a., J. T. Howell, and J. M. Linsdale. 1952. Plants of the Toiyabe Mountains area, Ne- vada. Wasmann J. Bio. 10:129-200. Lists 521 taxa; annotations of plant list include collection date, location, and number, geographic range, and elevation. Discu,sses climate, distribution of some major shmb and tree species, geographic af- finities of the flora. Compares flora to that of the Charleston Mountains. Includes photographs of plant communities, location map. New Mexico Bedker, E. J. 1966. A study of the flora of the Manzano Mountains. Albuquerque, NM: Univ. of New Mexico. Unpublished thesis. Lists .396 taxa; plant list annotated with collection locations. Discusses topography, climate, geology, soils and life zones; describes 17 study sites in the Manzano Moun- tains in terms of location, geological features, soils, plant communities, and dominant species. Provides keys to species, location map, and pho- tographs of plant communities. Compares flora to that of Mount Taylor, di.scu.s.ses species distri- bution in relation to environmental factors. March 1982 Bowers: Southwest Flora Bibliography 109 Campbell, R. S., and I. F. Campbell. 1938. Vegetation on gyp.sum soils of the Jornada Plain, New Mexi- co. Ecology 19:572-577. Lists 8 taxa based on 10 years of observation and collection. Discusses cli- mate, soil chemistry, and soil physical properties in relation to the depauperate flora and sparse vegetation. Capulin Mountain National Monument. 1980. Plant checklist: Capulin Mountain National Mon- ument, New Mexico. Capulin, NM: Capulin Mountain National Monument. Available from Capulin Mountain National Monument, Capulin, NM. Lists 172 taxa for .310 ha; plant list anno- tated with common names. Chaco Canyon National Monument. 1979. Checklist of plants for Chaco Canyon National Monument, New Mexico. Bloomfield, NM: Chaco Canyon National Monument. Available from Chaco Cul- ture National Historical Park, Star Rt. 4, Box 6500, Bloomfield, NM. Lists 247 taxa for 8705 ha; plant list annotated with common names and in- troduced species. The list is currently being up- dated by Anne Cully. Fletcher, R. A. 1972. A floristic assessment of the Datil Mountains. Albuquerque, NM: Univ. of New Mexico. Unpublished thesis. Lists 590 taxa for 82,902 ha; annotations include common names, phenology, elevational range, parent material, life form, relative abundance, geographical range, forage value, poisonous properties, and tax- onomic comments. Lists noteworthy collections, major collection sites, and birds and mammals observed in the area. Discusses land use, topogra- phy, geology, soils, and climate. Describes plant associations. Analyzes geographic affinities of the flora, compares flora to those of neighboring states, compares affinities and species richness of floras on different substrates. FosBERG, F. R. 1940. The aestival flora of the Mesilla Valley region. New Mexico. American Midland Naturalist 23:573-593. Lists 504 taxa; annotations of plant list include plant community and habi- tat. Describes plant communities and habitat; discusses topography and climate. Includes small- scale vegetation map. Gehlbach, F. R., B. H. Warnock, W. C. Martin, and H. K. Sharsmith. 1969. Vascular plants of Carls- bad Caverns National Park, New Mexico, and ad- jacent Guadalupe Mountains (New Mexi- co-Texas). Carlsbad, NM; Carlsbad Caverns National Park. Available from Carlsbad Caverns National Park, NM. Lists 640 taxa for 18,921 ha; annotations of plant list include common names, habit, and phenology. Hall, H. H., and S. Flowers. 1961. Vascular plants found in the Navajo Reservoir Basin, 1960, Colo- rado and New Mexico. Pages 47-90 in A. M. Woodbury, ed., Ecological studies of the flora and fauna of Navajo Reservoir, Colorado and New Mexico. Sah Lake City, UT: Univ. of Utah. Anthropological Paper No. 55. Lists 287 taxa for 6880 ha; annotations include common names, ele- vation, relative abundance, collector, and collec- tion location and date. HuTCHiNS, C. R. 1974. A flora of the White Mountains area, southern Lincoln and northern Otero coun- ties. New Mexico. Albuquerque, NM: C. R. Hut- chins. Lists 1686 taxa; location map provided; an- notations include synonymy, phenology, species de.scriptions, elevational range, habitat, and com- mon names. Provides keys to species and glo,ssary of botanical terms. Discusses topography and cli- mate. Lists subalpine taxa. Little, E. L., Jr., and R. S. Campbell. 1943. Flora of Jornada Experimental Range, New Mexico. Amer. Midi. Naturalist .30:626-670. Lists 528 taxa for 78,265 ha; annotations include geographic range of species, Raunkaier life form. Discusses history of botanical collection, geographic affi- nities of flora, distinctive species. Compares flo- ristic composition and plant cover of the Jornada plain with nearby mountain meadows; discusses forage value of various species. Lists weeds. Pro- vides location map and photographs of vegeta- tion types and plants. Mackay, H. a. 1970. A comparative floristic study of the Rio Hondo Canyon-Lake Fork- Wheeler Peak locale. New Mexico, and the Huerfano River- Bianco Peak locale, Colorado. Albuquerque, NM: Univ. of New Mexico. Unpublished di.ssertation. Lists 359 taxa for Rio Hondo locale, 467 taxa for Huerfano River locale; annotated with elevation- al ranges of each species. Discusses floristic dif- ferences between the two locales and describes vegetation zones. Collection sites are described in terms of topographic location, elevational range, and characteristic species. Describes geol- ogy and topography. Provides keys to species, vegetation maps at 1:63,000, photographs of veg- etation types. Manthey, G. T. 1977. A floristic analysis of the Sevilleta Wildlife Refuge and the Ladron Mountains. Al- buquerque, NM: Univ. of New Mexico. Unpub- lished thesis. Lists 728 taxa. Annotations of plant list include plant community, common name, phenology, and elevational range. Describes top- ography, geology, climate, soils, and land use. Discusses floristic affinities, community integrity of plant communities, geographic patterns of each community. Describes plant communities, listing major and common species, habitat, eleva- tional range, and local distribution of each. Lists taxa which terminate their distribution in or near the study area. Includes keys to species and a topographic map of the study area. Martin, W. C, C. R. Hutchins, and R. G. Woodmansee. 1971. A flora of the Sandia Moun- tains, New Mexico. Albuquerque, NM: Sandia Press. Lists 884 taxa; annotations include species descriptions, elevational range, common names, and synonymy. Includes keys to species and glos- sary of botanical terms. Discusses geology, vege- tation zones, and climate. Martin, W. C, and W. L. Wagner. 1974. Biological survey of Kirtland Air Force Base (East) [Micro- form]. Washington, D.C.: Government Printing Office; SAND-74-0393. Available from National Technical Information Service, Springfield, VA. 110 Great Basin Naturalist Vol. 42, No. 1 Lists 369 taxa for 19,430 ha; annotated with com- mon names, habitat, plant community. Describes topography, soils, climate; discusses biogeo- graphical influences and disturbance patterns. Etescribes plant communities. Includes checkHsts of amphibians, reptiles, mammals, and birds. OsBORN, N. L. 1962. The flora of Mount Taylor. Albu- querque, NM: Univ. of New Mexico. Unpub- lished thesis. Lists 301 taxa; includes keys to spe- cies. Discusses life zones; describes collection sites in terms of geographic location, elevation, life zone, geological features, dominant species. Includes checklist of all species collected at each site. Compares flora to that of other mountain ranges. 1966. A comparative floristic study of Mount Taylor and Redondo Peak, New Mexico. Albu- querque, NM: Univ. of New Mexico. Unpub- lished dissertation. Lists 231 taxa for Redondo Peak. Describes life zones, collection locations. Compares floras of the two peaks, listing species found on both. Compares populations of three species found on both peaks to detect possible evolutionary trends. Reviews paleobotany of the region. Includes keys to species. Riffle, N. L. 1973. The flora of Mount Sedgwick and vi- cinity. Albuquerque, NM: Univ. of New Mexico. Unpublished thesis. Lists 353 taxa; plant list an- notated with collection locations. Briefly dis- cusses geology and climate. Describes habitats of collection locations and lists dominant and inter- esting plants of each. Robertson, C. W. 1968. A study of the flora of Cochiti and Bland canyons of the Jemez Mountains. Al- buquerque, NM: Univ. of New Mexico. Unpub- lished thesis. Lists 373 taxa. Discusses topogra- phy, soils, geology, and climate. Describes plant communities of each elevational zone. Compares floras of Cochiti and Bland canyons. Provides keys to species; includes topographic map and photographs of topography and vegetation. Plant checklist is annotated with collection locations. ScHAFFNER, E. R. 1948. Flora of the White Sands Na- tional Monument of New Mexico. Albuquerque, NM: Univ. of New Mexico. Unpublished thesis. Lists 110 taxa for 56,757 ha; annotations include common names, local distribution, relative abun- dance, ethnobotanical notes, palatability for live- stock, synonymy, and identification notes. In- cludes keys to species, photographs of vegetation, and topography. Discusses geology, topography, climate, collection history, and adaptation of plants to dunes. Lists gypsum indicator plants. Tatschl, a. K. 1966. A floristic study of the San Pedro Parks Wild Area, Rio Arriba County, New Mexi- co. Albuquerque, NM: Univ. of New Mexico. Un- published thesis. Lists 288 taxa. Briefly describes topography, soils, and climate. Discusses life zones, lists representative species. Describes nine major collecting locations in terms of elevational range, dominant plants, habitat. Provides keys to species. Includes location and physiographic maps, photographs of plant communities. Plant checklist annotated with collection locations. Van Devender, T. R., and B. L. Everitt. 1977. The lat- est Pleistocene and Recent vegetation of the Bishop's Cap, south central New Mexico. South- western Naturalist 22:337-352. Lists 118 taxa. Discusses climate, packrat ecology, modern and fossil floras in relation to climatic change. Lists 39 plant species found as macrofossils in packrat middens dating from 10,500 B.P. Wagner, W. L. 1973. Floristic affinities of Animas Mountain, southwestern New Mexico. Albu- querque, NM: Univ. of New Mexico. Unpub- lished thesis. Lists 718 taxa for 96,000 ha; annota- tions of plant list include life form, habitat, relative abundance, geographic range, plant com- munity, synonymy, taxonomic comments and en- demic species. Describes climate and topogra- phy. Discusses factors in plant distribution and dispersal, evolution of the flora, and distribution- al patterns of the various life forms. Defines Northern Sierra Madre Biotic Province, analyzes geographic affinities of the flora and the plant communities. Describes collection sites and plant communities in terms of dominant perennials and characteristic annuals, elevational range, ecologi- cal factors and topographic location. Wynhoff, J. T., E. Lehto, and D. J. Pinkava. 1976. Vegetation of Cutler Canyon and vicinity, San Juan County, New Mexico. Washington, D.C.: U.S. Dept. of the Interior, Bureau of Land Man- agement. Lists 436 taxa; annotations include rela- tive abundance, common name, habitat. Includes photographs of habitat. Fide Donald J. Pinkava, Arizona State University. Utah Allred, K. W. 1975. Timpanogos flora. Provo, UT: Brig- ham Young Univ. Thesis. Lists 684 taxa for 15,544 ha; annotations of plant list include habi- tat, phenology, elevational range, and collector and collection number. Discusses history of bo- tanical collection. Describes topography, climate, vegetation zones. Lists dominant and character- istic species of each vegetation zone. Includes keys to species. Arnow, L. a. 1971. Vascular flora of Red Butte Canyon, Salt Lake County, Utah. Salt Lake City, UT: Univ. of Utah. Unpublished thesis. Lists 511 taxa for 2228 ha; annotations include species descrip- tions, common names, phenology, habitat, tax- onomic comments, and plant associations. Briefly discusses topography, climate, geology, and his- tory of botanical collection. Includes keys to spe- cies and glossary of botanical terms. Arnow, L. A., B. J. Albee, and A. M. Wyckoff. 1980. Flora of the central Wasatch Front, Utah. Salt Lake City, UT: Univ. of Utah Printing Service. Available from Univ. of Utah Bookstore, Salt Lake City, UT. Lists 1139 taxa for 260,000 ha. Briefly discusses vegetation of elevational zones, climate, and topography. Annotations include synonymy, common names, species descriptions, phenology, habitat, and elevational range. Pro- vides keys to species and an illustrated glossary of botanical terms. March 1982 Bowers: Southwest Flora Bibliography 111 Buchanan, H., and G. T. Nebeker. 1971. Checklist of the higher plants of Bryce Canyon National Park, Utah. Ogden, UT: Weber State College Printing Dept. Lists 324 taxa. Describes and illustrates habitats of Bryce Canyon. Plant list annotated with common names. Fide Donald J. Pinkava, Arizona State University. Castle, E. D. 1954. The succession of vegetation on a southern Utah sand dune. Provo, UT: Brigham Young Univ. Unpublished thesis. Lists 65 taxa; annotations of plant list include plants collected near the dunes. Discusses physiography, climate, and dune plant morphology. Describes vegeta- tion on active and stable dunes. Discusses plant succession, limiting factors for plants on sand. In- cludes photographs of vegetation and topography. Flowers, S., H. H. Hall, and G. T. Groves. 1959. An- notated list of plants found in Flaming Gorge Reservoir Basin. Pages 49-98 in A. M. Woodbury, ed., Ecological studies of the flora and fauna of Flaming Gorge Reservoir Basin, Utah and Wyoming. Salt Lake City, UT: Univ. of Utah; 1960. Anthropological Paper No. 48. Lists 448 taxa for river length of 142 km; annotations in- clude relative abundance, habitat, elevational range, collection locations, and collector. Graham, E. H. 1937. Botanical studies in the Uinta Ba- sin of Utah and Colorado. Annals of the Carnegie Museum, Pittsburgh 26. Lists 1104 taxa for 3,108,808 ha; annotations include common names, relative abundance, habitat, collection lo- cation, elevation, vegetation zone, phenology and cx)llector, collection date and number. Discusses history of exploration and settlement in Utah, history of botanical collection. Describes topogra- phy, climate, geology; vegetation zones, com- paring them with those of surrounding areas; plant communities, hsting dominant and second- ary species of each. Eighty-nine collection loca- tions are described in terms of location, plant community, elevation, and substrate. Discusses geographic affinities of flora; lists range exten- sions, new varieties, and species. Harrison, B. F., S. L. Welsh, and G. Moore. 1964. Plants of Arches National Monument. Brigham Young Univ. Sci. Bull., Biol. Ser. 5(1): 1-23. Lists .322 taxa for 29,696 ha; annotations include col- lection location and collector, relative abun- dance, habitat, local distribution, and common names. Briefly discusses climate, geology, and habitat. Lists endemics, discusses introduced species. Higgins, L. C. 1967. A flora of the Beaver Dam Moun- tains. Provo, UT: Brigham Young Univ. Unpub- hshed thesis. Lists 667 taxa; annotations of pla rt list include geographic range, relative abun- dance, elevational range, collector, and collection number. Briefly describes geology, major plant communities, topography. Discusses history of botanical collection. Includes keys to species. Lists range extensions, endemic species. Holmgren, A. H. 1948. Handbook of the vascular plants of the northern Wasatch. 3d ed. Palo Alto, CA: National Press. Lists 1266 taxa, including many cultivated plants. Annotations include common names, habitat, local distribution, synonymy. In- cludes keys to species and glossary of botanical terms. Lewis, M. E. 1973. Wheeler Peak area species list. Og- den, UT: U.S. Forest Service. Available from In- termountain Region, U.S. Forest Service, Ogden, UT. Lists 409 taxa; annotations of plant list in- clude plant community, habitat, and relative abundance. McMillan, C. 1948. A taxonomic and ecological study of the flora of the Deep Creek Mountains of cen- tral western Utah. Salt Lake City, UT: Univ. of Utah. Unpublished thesis. Lists 431 taxa; annota- tions of plant list include relative abundance, lo- cal distribution, and habitat. Describes phys- iography, climate, vegetation zones. Discusses history of botanical collection, effects of glacia- tion, phytogeography. Includes photographs of vegetation types. Meyer, S. E. 1976. Annotated checklist of the vascular plants of Washington County, Utah. Las Vegas, NV: Univ. of Nevada. Unpublished thesis. Lists 1207 taxa for 621,762 ha; annotations of plant list include habitat, local distribution, floristic com- ponent, synonymy, community type, collector, and collection date and number. Reviews history of botanical collection. Discusses physiography and geology; divides county into six topographic units and discusses the elevational range, accessi- bility, geology, and watersheds of each. Describes climate and soils and discusses climatic and eda- phic factors affecting plant distribution. De- scribes community types and plant associations; discusses impact ol^ grazing and land use on vege- tation. Defines floristic components. Meyer, S. E. 1980. Capitol Reef National Park, revised working checklist. Torrey, UT: Capitol Reef Na- tional Park. Available from Capitol Reef National Park, Torrey, UT. Lists 472 taxa; plant list anno- tated with common names. Nebeker, G. T. 1975. Manual of the flora of the East Tintic Mountains, Utah. Provo, UT: Brigham Young Univ. Unpublished thesis. Lists 211 taxa; annotations include habitat, geographic range, collection number. Briefly discusses topography, geology, plant communities. Includes keys to species. Nelson, R. A. 1976. Plants of Zion National Park. Springdale, UT: Zion Natural History Associ- ation. Available from Zion National Park, Spring- dale, UT. Lists 704 taxa for 59,307 ha; annota- tions include species descriptions, local distribution, relative abundance, habitat, and common names. Describes vegetation zones, dis- cusses phenology. Includes keys to species, glos- sary of botanical terms, photographs of plant communities, and line drawings of many plants. Preece, S. J., Jr. 1950. Floristic and ecological features of the Raft River Mountains of northwestern Utah. Salt Lake City, UT: Univ. of Utah. Unpub- lished thesis. Lists 341 taxa; annotations include relative abundance, habitat, elevational range, collector, and collection location and number. 112 Great Basin Naturalist Vol. 42, No. 1 Describes topography and geology. Discusses his- tory of botanical collection; vegetation types, correlating vegetation with substrate, climate, and grazing; and floristic elements and relation of flora to paleoclimate. Includes photographs of vegetation. Welsh, S. L. 1957. An ecological survey of the vegeta- tion of the Dinosaur National Monument, Utah. Provo, UT: Brigham Young Univ. Unpublished thesis. Lists 360 taxa for 30,352 ha; annotations of plant list include geologic formation, habitat, as- sociated species, elevation, collector, and collec- tion location and number. Describes geology, to- pography, and climate. Describes plant communities on different geological formations. Lists major species of each plant community, dis- cusses trends in .succession and edaphic factors in plant distribution. Includes photographs of geo- morphological features, vegetation map, topo- graphic map, and geological map. Welsh, S. L., and G. Moore. 1968. Plants of Natural Bridges National Monument. Proc. Utah Acad. Sci., Arts, and Letters 45:220-248. Lists 224 taxa for 3148 ha; annotations of plant list include common names, relative abundance, habitat, col- lector, and collection location, date, and number. Describes plant communities in relation to topog- raphy and habitat. Literature Cited Bowers, J. E. 1981. Local floras of Arizona: an anno- tated bibliography. Madrono. 28:193-209. Christensen, E. M. 1967a. Bibliography of Utah botany and wildland conservation. Brigham Young Univ. Sci. Bull., Biol. Ser. 9(1): 1-136. 1967b. Bibliography of Utah botany and wild- land conservation. No. II. Proc. Utah Acad. Sci., Arts, and Lett. 44:545-566. Cronquist, a., a. H. Holmgren, N. H. Holmgren, J. L. Reveal, and P. Holmgren. 1972. Intermountain flora: vascular plants of the Intermountain Re- gion, USA. Vol. 1. Hafner, New York. 1977. Intermountain flora. Vol. 6. Columbia Univ. Press, New York. EwAN, J. 1936. Bibliography of the botany of Arizona. American Midland Naturalist 17:430-454. Harrington, H. D. 1954. Manual of the plants of Colo- rado. 2d ed. Sage Books, Denver, Colorado. Hoffman, G. R., and G. J. Tomlinson. 1966. A bibliog- raphy of vegetational studies of Colorado. South- western Naturalist 11:223-237. Holmgren, A. H., and J. L. Reveal. 1966. Checklist of the vascular plants of the Intermountain Region. U.S. Forest Service, Intermountain Forest and Range Experiment Station, Ogden, Utah. Re- search Paper No. 32. Jaeger, E. C. 1941. Desert wild flowers. Rev. ed. Stan- ford Univ. Press, Stanford, California. Kearney, T. H., and R. H. Peebles. 1942. Flowering plants and ferns of Arizona. U.S. Dept. of Agri- culture, Washington, D.C. Miscellaneous Pub- lication No. 423. 1960. Arizona flora. Univ. of California Press, Berkeley, California. Martin, W. C, and C. R. Hutchins. 1981. Flora of New Mexico. Lubrecht and Cramer, Monticello, New York. ScHMUTz, E. M. 1978. Cla.ssified bibliography on native plants of Arizona. Univ. of Arizona Press, Tucson, Arizona. Shreve, F., and I. L. Wiggins. 1964. Vegetation and flora of the Sonoran Desert. Stanford Univ. Press, Stanford, California. TiDESTROM, I. 1925. Flora of Utah and Nevada. U.S. Nat. Herb. Contrib. 25. TiDESTROM, I., AND T. KiTTELL. 1941. A flora of Arizona and New Mexico. Catholic Univ. of America Press, Washington, D.C. U.S. Fish and Wildlife Service. 1977a. Annotated bib- liography of natural resources information: north- western New Mexico. FWS/OBS-77/33. 1977b. Annotated bibliography of natural re- sources information: southern Utah. FWS/OBS- 77/34. 1977c. .Annotated bibliography of natural re- sources information: northwestern Colorado. FWS/OBS-77/35. Welsh, S. L., and G. Moore. 1973. Utah plants; tracheophvta. Brigham Young Univ. Press, Provo Utah. Acknowledgments I thank Dr. Donald Pinkava, Dr. Stanley Welsh, Dr. William Martin, and Dr. Arthur Holmgren for reviewing the manuscript and suggesting additional floras. I also thank Dr. Elizabeth Neese, Dr. Lois Arnow, Dr. Kim- ball Harper, and Dr. Beverly Albee, who pro- vided several floras. The interlibrary loan personnel at the University of Arizona Li- brary obtained many books and dieses for me, and without their invaluable assistance I could not have compiled this bibliography. THE RELATION BETWEEN SPECIES NUMBERS AND ISLAND CHARACTERISTICS FOR HABITAT ISLANDS IN A VOLCANIC LANDSCAPE Steven H. Carter-Lovejoy' Abstract.- Within the Craters of the Moon Lava Flow in southeastern Idaho are kipukas, islands of sagebrush habitat isolated by relatively barren lava. In 1979 I counted the number of species of plants, small mammals, and reptiles for a series of kipukas. The degree of isolation of a kipuka was not related to numbers of species found there; thus, these data do not support the equilibrium theory of island biogeography. For most of the study organisms lava is not a significant barrier to dispersal. Larger kipukas support larger numbers of species. For plants this relationship likely is a result of the increased topographic variety of larger kipukas, whereas for small mammal species minimum area requirements for maintaining populations are met only by larger kipukas. More distant kipukas showed in- creased density of small mammals\ possibly as a result of reduced predation. Patterns of distribution in this ecosys- tem are best explained, not by any all-inclusive community mechanism, but through the agglomerative contributions of a variety of population and community fimctions. The Snake River Plain in Idaho is a forma- tion built up, over millions of years, of wind- blown loess deposits alternating with layers of volcanic materials from localized erup- tions (Greeley and King 1975). The present manifestation of this geologic history is a complex and heterogeneous combination of soils and surfaces, ranging from well-drained loess soils dozens of feet deep to bare lava rock surfaces. The distribution of vegetation is dictated by this physiographic hetero- geneity; species found growing on a lava sur- face differ from those found in a deep crevice in the lava surface, and both sets of species differ from species fovmd on deep soil that might occur a few meters away. The distribu- tion of animals also reflects the heterogeneity of their environment, both physical and flor- al. The impact of edaphic complexity on plant and animal distributions is enhanced by unevenly distributed sources of dispersal. Kipuka is a name given to an area of older land surrounded by but not buried by more recent lava flows. Within the most recent lava flows on the Snake River Plain, kipukas present striking contrasts in landscapes. The approximately 150,000 ha lava surface of the Craters of the Moon Flow is about 2000-2500 years old (Prinz 1970)-recent enough, in the cold, arid climate of south- eastern Idaho, that little soil has accumulated and little vegetative colonization of the lava has taken place (Eggler 1941). Superimposed on this barren landscape are hundreds of ki- pukas; although these kipukas are diverse in age, edaphic and topographic characteristics, and isolation, all are well vegetated. In the summer of 1979, I studied the distri- bution of plants, small mammals, and reptiles on a number of kipukas within the Craters of the Moon Flow in southeastern Idaho. This paper presents data relevant to the question. To what extent does the islandlike nature of kipukas shape the observed floral and faunal distributions? Methods Study sites were located on 14 kipukas sit- uated near the eastern edge of the Craters of the Moon Flow, 4-21 km south of Arco, Idaho (Fig. 1). The area is characterized by a continental climate, with cold winters and hot summers; mean annual precipitation, fall- ing mostly as snow, is only about 25 cm, but it fluctuates considerably from year to year (Lovejoy 1980). The region lies within Kuch- ler's (1975) sagebrush-steppe vegetation zone; ■Department of Biology, Idaho State University, Pocatello, Idaho 83209. Current address; Department of Biological Science, Florida State University, Tal- lahassee, Florida 32306. 113 114 Great Basin Naturalist Vol. 42, No. 1 dominant shrubs are Artemisia tridentata (big sagebrush) and A. tripartita (threetip sage- brush), and dominant grasses are Agropyron spicatum (bluebunch wheatgrass) and Stipa spp. (needle-and-thread), but the importance of these and of other plant species varies broadly from kipuka to kipuka and even within kipukas. The study kipukas range in size from 0.16 to 3.6 ha and are separated from the nearest off-lava vegetation by 70-1800 m. From one to four study plots were established on each kipuka. An effort was made to represent ade- quately the subjectively perceived environ- mental diversity on each kipuka. Slope and aspect of each plot were measured and soil character measures made. Vegetation was sampled by canopy-cover estimation (Daubenmire 1959, Hanley 1976). Percent cover of all perennial plants was esti- mated for 30 0.2 by 0.5 m quadrats situated LLO National Monument 40 km Fig. 1. a, The Snake River Plain in southern Idaho, shown with hatchmarks. b, Recent lava flows in south- eastern Idaho. The study kipukas are located within the stippled area. systematically within each study plot. Species recorded during the systematic samplings were added to the comprehensive species list developed by a walking survey for each ki- puka. The total plant species pool of the study area is small; fewer than 80 species on lava or kipukas were encountered during the field study, so the species lists are rather complete. In each plot, 18 trapping stations were lo- cated 10 m apart in a 6 by 3 grid, and three Museum Special snap traps and one McGill rat trap were set at each station. Bait was peanut butter and rolled oats; after a three- day prebait period, traps were checked and reset for three or four nights. Additional traps were located in likely areas on some kipukas and in some lava areas between kipukas and the mainland. Trapping was done within a six-week period (May-June) to minimize the potential for seasonal differences in captures among kipukas (Smith et al. 1975, Johnson 1977). Lizard populations were assessed on six kipukas with unbaited pitfall traps. Liz- ards captured were marked and released, but marked lizards were never recaptured. Spe- cies lists of reptiles and of small mammals were augmented by personal sightings. Results The number of plant species on the ki- pukas ranged from 25 to 49. The number of plants on a kipuka is significantly related to the size of the kipuka (Fig. 2), but not to the degree of isolation of the kipuka (Table 1). The prominent vegetative patterns were re- lated to edaphic and topographic character- istics; these patterns were present within as well as among kipukas (Lovejoy 1980). A companion paper presents the results of ordi- nations used to explore these patterns (Carter-Lovejoy, in preparation). Reliable assessments were made of the presence of eight species of small mammals and two lizard species on appropriately trapped kipukas. The number of lizard spe- cies on the six pitfall-trapped kipukas ranged from zero to two; species captured were Scehporus graciosus (sagebrush lizard) and Eumeces skiltonianus (western skink). The number of small mammals on a kipuka ranged from three to seven. Four species March 1982 Carter-Love joy: Habitat Islands 115 were widely distributed. Peromyscus manicu- latus (white-footed deer mouse) was captured wherever traps were set, and Etitamias mini- mus (least chipmunk) was caught in most lo- calities. As a result of their larger sizes, Neo- toma cinerea (bushy-tailed woodrat) and Sylvilagus nuttalli (Nuttall's cottontail) were captured less often; however, S. nuttalli indi- viduals were observed on every kipuka, and characteristic urine markings of N. cinerea were also ubiquitous. Four other species were captured less often: Perognathus parvus (Great Basin pocket mouse) was captured on two kipukas, Microtus montanus (montane vole) on four, Reithrodontomys megalotis (western harvest mouse) on one, and Sperrno- philus townsendii (Townsend's ground squir- rel) was caught on three kipukas and sighted on a fourth. As with the plant species number, the number of small mammal species on a kipuka is positively correlated with size of the ki- puka (Fig. 3). The number of mammal spe- cies on a kipuka is not correlated with isola- tion of the kipuka, however (Table 1). I.8n Fig. 2. a kipuka P < 0.05. —I 1 r 0.4 0.£ Species-area as a fiinct 1.2 1.6 2.0 KIPUKA AREA (ha) 2.4 Discussion The equilibrium theory of island biogeo- graphy (MacArthur and Wilson 1967) postu- lates that the number of species on an island is determined by a dynamic equilibrium be- tween immigration rate and extinction rate. The theory further maintains that two phys- ical measures, island isolation and island size, successfully predict species immigration and extinction rates. Since its conception this the- ory has been widely accepted and applied by ecologists; it has recently been seriously chal- lenged, however, for its feeble empirical sup- port (Gilbert 1980). The equilibrium theory has been applied to several types of habitat island, often with inconsistent and poorly in- terpreted results (see Gilbert 1980). In this study the theory-derived prediction that more isolated kipukas have fewer species is not borne out. Application of the MacArthur-Wilson species-isolation model has rarely been made to plants (Simberloff 1974, but see Crowe 1979). The persistence of individuals of pe- rennial species is long relative to immigration rates. Only in systems with very broad bar- riers to plant dispersal or with short-lived plants dominant would the presumed isola- tion effect be important. Certainly Snake River Plain vegetation and the system of lava Table 1. Small mammal and plant species numbers for kipukas that are grouped into size classes and ar- ranged according to degree of isolation. Distance No. No. Kipuka Size to M.L.^ mammal plant name (ha) (m) specie^ species 2 .45 70 3 25 7 .16 350^ 5 27 4 .40 400 3 39 13 .69 550 4 28 14 .57 560 4 31 9 .57 1490 4 27 1 1.4 160 5 .30 5 1.1 410 4 41 15<= 2.2 670 4 31 10 .97 1800 5 28 6 3.1 220 7 49 8 3.6 500 6 37 15^- 2.2 670 4 31 16 ,3.2 1270 6 31 effect; number of plant species on ion of the size of the kipuka. ^Distance to mainland: shortest direct-line distance across lava. ''Number of small mammal species: numbers include Neotoma cinerea and Sylvilagus nuttalli. presumed present on every kipuka. "^Kipuka 15 is placed in two size classes because of its intermediate size. °Only 70 m from kipuka 6. 116 Great Basin Naturalist Vol. 42, No. 1 barriers do not fit these models. Immigration of plant propagules into kipukas probably differs little from the cross-seeding that oc- curs among the heterogeneous habitats of the Snake River Plain. If an isolation effect did occur, one would be more likely to observe it with small mam- mals; I did not observe an isolation effect. It is worthwhile to evaluate to what degree lava acts as a barrier to small mammal dis- persal. Eutamias minimus, P. maniculatiis, S. nuttalli, and N. cinerea were captured in traps set on lava. Scat and urine markings of the latter two species were frequently sighted up to several kilometers from any kipuka or mainland, and it is quite possible that the lat- ter three nest and breed on lava; certainly N. cinerea does. Although the small amount of vegetation on the lava may limit population sizes of these four species on lava, it appar- ently does not restrict their ability to disperse across it. If kipukas are not functional isolates but are, at best, the most favorable patches with- in a matrix of suitable habitat, then extinction on a kipuka is unlikely; indeed, in this con- text "extinction" has little meaning. Since in- dividuals of these four readily dispersing spe- cies made up over 96 percent of all small 1.0 2.0 3.0 KIPUKA AREA (ha) 4.0 Fig. .3. Species-area effect: number of small mammal species on a kipuka as a fimction of the size of the ki- puka. P < 0.01. mammal captures, examination of aggregate small mammal data should obviously reveal no isolation effect. It is much less likely that the four other small mammal species trapped here actually reside on lava. One M. montanus individual was sighted one night on lava less than 50 m from both a kipuka and the mainland; in no other instance were any of these species sighted or captured on lava. On the other hand, they were not very commonly sighted on kipukas either. Only 24 individuals of these four species were captured. Given the high trapping intensity on kipukas, it is pos- sible to assume that individuals representing the sole capture of their species on any ki- puka are immigrants to the kipuka, not mem- bers of established populations. There were 11 encounters of these four species during the study; in only 4 cases were we sampling a population (conservatively defined as two or more individuals). The other 7 encounters were of single individuals, despite exhaustive trapping of the habitats in which these indi- viduals were found. Isolation seemed to play no role in the presence of the two lizard species found on the kipukas either. On the other hand, on no kipuka did we find the diurnal and con- spicuous Phrynosoma doiiglassi (desert short- horned lizard), even though it was frequently sighted on mainland; its total absence on ki- pukas suggests that lava is more of a barrier to the dispersal of this slow-moving lizard than it is to the dispersal of the two quicker species. One interesting and unexpected isolation effect does exist: the density of animals is sig- nificantly and positively correlated with iso- lation (Fig. 4). Common explanations for so- called "density compensation" (MacArthur 1972) are not appropriate in light of the fact that the density measure, individuals per 100 trap nights, reflects changes in the density of the common species, to which lava is not a barrier. One possible mechanism for in- creased density on isolated kipukas is reduced predation. Densities of prey for common pre- dators in the area are higher on kipukas than they are on nearby lava, because of the rela- tive abundance of vegetation on kipukas. As a result of this concentration of prey, pre- dators are likely to focus their attention on March 1982 Carter-Lovejoy: Habitat Islands 117 kipukas. Predation pressure is intermittent, however, particularly on smaller kipukas. Small kipukas (less than 3 ha) are probably not large enough to support a snake popu- lation (Lovejoy 1980). We rarely observed birds of prey over the study area, perhaps be- cause of the lack of roosts. Although scat finds indicated that coyotes were occasional visitors to all study kipukas, none of the ki- pukas is large enough to support a resident coyote. So, between times of possibly intense predation, kipukas may enjoy periods of total release from predation pressure. During these lulls, small mammal populations may be able to build up to densities that greatly exceed densities on less isolated kipukas, where pre- dators may forage more frequently. Certainly this density-isolation phenomenon deserves further study. Connor and McCoy (1979) discuss three mechanisms that can explain an observed species-area effect. The "habitat-diversity" hypothesis (Williams 1964) postulates that, as increasingly larger areas are sampled, new habitats with associated new species are en- countered. According to the "area-per se" hypothesis (Preston 1960, MacArthur and Wilson 1967), population sizes are propor- tionately smaller on smaller isolates, implying increased probability of species extinctions. An alternative "passive sampling" hypothesis (Connor and McCoy 1979) holds that species number is controlled by passive sampling from a species pool, so that larger areas inter- cept larger samples than smaller areas. Con- nor and McCoy consider this a good "null" hypothesis. The area-per se hypothesis is associated with the equilibrium island biogeography model; the implication is that an ecologically significant rate of species extinction results in a tiu-nover of species. A number of studies (Gilbert 1980:214, cites 23) purport to sup- port the equilibrium model by demonstrating species-area relationships among island groupings, despite the fact that an observa- tion of a species-area relationship made in a temporally limited study cannot demonstrate species turnover and thus cannot distinguish between the three causal hypotheses above. Specific manipulative experimentation is necessary to distinguish among these hypoth- eses; unfortunately, such experimentation is virtually nonexistent. Without such experi- mentation, one can only speculate about causes for the observed relationship. Data from this study could be used to sup- port all three hypotheses, without refuting any of them. Certainly environmental hetero- geneity increases with increasing kipuka size, because larger kipukas have more topograph- ic variation. Support for the null hypothesis can be noted by looking strictly at the num- bers of small mammal immigrants on kipukas of dfferent sizes (Fig. 5). These immigrants have presumably reached the kipukas on which they were found by chance; never- theless, they show a species-area effect, in- dicating the kipukas are passively sampling small mammal dispersers. The area-per se hypothesis also appears to have some application to these data. Perhaps the most intrinsically acceptable explanation for the species-area relationship lies with the population ecology of the different species contributing to species number. Equilibrium theory's contention that extinction rates de- cline with increasing area is by no means cer- tain; on the other hand, decreasing area will at some point result in the extinction of any 2ln 2 4 6 8 10 12 14 16 18 20 KIPUKA ISOLATION (xlOOm) Fig. 4. Density-isolation effect: small mammal density (number of individuals captured for every 100 nights of trapping) as a function of kipuka isolation (straight-line distance to nearest mainland). P < 0.05. 118 Great Basin Naturalist Vol. 42, No. 1 particular population. Thus, there must exist some threshold size for each species, below which the probability of extinction of a pop- ulation of this species is one (Shaffer 1981). Of the small mammal species observed on ki- pukas for which lava poses an impediment to dispersal, three species appear to have had established populations on at least one ki- puka: P. parvus, S. townsendii, and M. mon- tanus. None of these populations was on a ki- puka less than three ha in size (Fig. 5). The species involved did not appear to be re- stricted to a particular type of habitat that was unavailable on smaller kipukas, nor did the increase in environmental heterogeneity on kipukas of larger size appear to be impor- tant to the individual species. Instead, it is possible that these species need an area of at least three hectares in this ecosystem to sup- port a large enough number of individuals to maintain a population. Perhaps those species that did not have populations on any of the study kipukas have minimum area require- ments larger than three ha. 40 KIPUKA AREA (ha) Fig. 5. Species-area effect: number of rare small mammal species on a kipuka as a function of kipuka size. Dots indicate number of species with one or more individuals; X's indicate number of species witfi two or more individuals. Species considered rare are P. parvus, S. townsendii. M. montaniis, and R. memlotis P< 0.001. The minimum size requirements of many species relate to energetics and the available food resource (McNab 1963, Harestad and Bunnell 1979). Species vary in the amount of food required for living and reproducing: larger animals require more food, as do meta- bolically more active animals. This might ac- count for the absence of Lepus californicus (blacktail jackrabbit) and Dipodomys ordii (Ord's kangaroo rat) from all study kipukas, even though both species were common on nearby off-lava sites. Animals at higher posi- tions in community trophic structures require more cumulative biomass to stay alive, and populations of such animals will require larger areas in which individuals can forage for food. The absence of the carnivorous (Onychomijs leucogaster (grasshopper mouse) might be explained by this reasoning. Snakes— Co/t//7er constrictor (racer), Crotalus viridus (Great Basin rattlesnake), and Pi- tuophis melanoleucus (Great Basin gopher snake)— were sighted only on those kipukas or kipuka archipelagos that are greater than three ha in size. Social behavior is another factor that plays a role in the determination of minimum area requirements, although social behavior is, of course, largely an evolutionary response to energetic considerations. The home range of an animal is an expression of both the social and energetic factors, so that home range sizes, when known, are a rough indication of relative minimum size requirements for small mammals. Literature values for the home range sizes of two mainland species not found on kipukas (1.3-1.4 ha for O. leucogaster, 2.7-4.6 ha for D. ordii [French et al. 1975]) are considerably larger than those for kipuka- dwelling P. parvus (.05-.4 ha [O'Farrell et al. 1975]). Of course, alternative explanations exist for the absence of the small mammal species that were not found. There might be a lack of suitable habitat on kipukas for some species; for instance, D. ordii is generally associated with sandy soils, and only one trapped kipuka met this criterion. If the missing species was absent as well from the mainland immedi- ately adjacent to the lava flow, there would be no ready source of immigrants to colonize kipukas. Lava is probably a very effective. March 1982 Carter-Lovejoy: Habitat Islands 119 perhaps impenetrable barrier to some spe- cies. Certainly the observed distribution of small mammal species can be explained, not through any grand mechanism, but through the agglomerative contributions of a variety of population and community functions. The best imderstanding of patterns of distribution in the kipuka-lava ecosystem will primarily be elaborated, in the future, by autecological research. Acknowledgments I thank Janice Carter-Lovejoy for her help in the field, Barry Keller for traps and advice, and Dan Simberloff for comments on this pa- per. I am grateful for the guidance of Dr. Jay E. Anderson throughout this project. This work was completed as part of a contract from the Bm-eau of Land Management to Idaho State University. Literature Cited Connor, E. F., and E. D. McCoy. 1979. The statistics and biology of the species-area relationship. Amer. Nat. 113:791-a33. Crowe, T. M. 1979. Lots of weeds; insular phytogeo- graphv of vacant urban lots. J. Biogeography 6:169-182. Daubenmire, R. F. 1959. Canopy coverage method of vegetation analysis. Northwest Sci. 33:43-64. Eggler, W. a. 1941. Primary succession on volcanic de- posits in southern Idaho. Ecol. Monogr. 11:277-298. French, N. R., D. M. Stoddart, and B. Bobek. 1975. Patterns of demography in small mammal popu- lations. Pages 7.3-102 in F. B. Golley, K. Petruse- wicz, and L. Ryszkowski, eds.. Small mammals: their productivity and population dynamics. Cambridge Univ. Press, New York. Gilbert, F. S. 1980. The eq\iilibrium theory of island biogeographv: fact or fiction? J. Biogeography 7:209-2.35. Greeley, R., and J. S. King. 1975. Geologic field guide to the Quaternary volcanics of the south central Snake River plain, Idaho. Idaho Bureau of Mines and Geology, Moscow. Hanley, T. a. 1978. A comparison of the line- interception and quadrat estimation methods of determining shrub canopy coverage. J. Range Manage. 31:60-62. Harestad, a. S., and F. L. Bunnell. 1979. Home range and body weight— a reevaluation. Ecology 60:389-402. Johnson, W. C. 1977. Examination of censusing tech- niques for small mammals in a high desert ecosys- tem. Unpublished thesis. Idaho State Univ., Pocatello. Kuchler, a. W. 1975. Potential natural vegetation of the conterminous United States (map). American Geographical Society, New York. Lovejoy, S. H. 1980. Patterns in the distribution of plants and animals on lava flows and kipukas in southeastern Idaho. Unpublished thesis. Idaho State Univ., Pocatello. Mac Arthur, R. H. 1972. Geographical ecology. Harper and Row, New York. MacArthur, R. H., and E. O. Wilson. 1967. The theo- ry of island biogeography. Princeton Univ. Press, Princeton, New Jersey. McNab, R. K. 1963. Bioenergetics and the determina- tion of home range size. Amer. Nat. 97:133-140. O'Farrell, T. p., R. J. Olson, R. O. Gilbert, and J. D. Hedlund. 1975. A population of Great Basin pocket mice, Perognathus parvus, in the shrub- steppe of south central Washington. Ecol. Mon- ogr. 45:1-28. Preston, F. W. 1962. The canonical distribution of com- monness and rarity. Ecology 43:185-215, 410-432. Prinz, M. 1970. Idaho rift system. Snake River plain, Idaho. Geol. Soc. of Amer. Bull. 81:941-948. Shaffer, M. L. 1981. Minimum population sizes for spe- cies conservation. Bioscience 31:131-134. Simberloff, D. S. 1974. Equilibrium theory of island biogeography and ecology. Ann. Rev. Ecol. Syst. 5:161-182. Smith, M. H., R. H. Gardner, J. B. Gentry, D. W. Kaufman, and M. H. O'Farrell. 1975. Density estimations of small mammal populations. Pages 25-54 in F. B. Golley, K. Petrusewicz, and L. Ryszkowski, eds., Small mammals: their produc- tivity and population dynamics. Cambridge Univ. Press, New York. Williams, C. B. 1964. Patterns in the balance of nature. Academic Press, London. TAXONOMIC STUDIES OF DWARF MISTLETOES (ARCEUTHOBIUM SPP ) PARASITIZING PINUS STROBIFORMIS Robert L. Mathiasen' Abstract.- Analysis of morphological characters oi Arceuthobium apachecum and A. blumeri indicates there are several geographically consistent differences between these taxa, which supports their current classification at the specihc level. Shoot height and perianth lobe number exhibited considerable geographic variation, and some mor- phological characters examined were continuous. Peak flowermg and seed dispersal periods for these species differed slightly Altitudinal, seasonal, and latitudinal variations in flowering and seed dispersal were detected and may be re- sponsible for the differences in phenology between these taxa. Dwarf mistletoes (Arceuthobium spp.) are the most serious disease agents of conifers in the southwestern United States. Many of the North American species of Arceuthobium were described by Engehnann in the 1800s (Gray 1850, Watson 1880). Gill (1935) pre- pared the first comprehensive monograph of the genus, in which he reduced several pre- viously recognized fall-flowering species to host forms of A. campylopodum Engelm. Gill designated forms of A. campylopodum ex- clusively on the basis of host relationships be- cause he recognized few morphological dif- ferences between these taxa. In Gill's system A. campylopodum Engelm. forma blumeri (Engelm.) Gill encompasses the dwarf mis- tletoes that parasitize sugar pine {Pinus lam- hertiana Dougl.) and western white pine {Pinus monticola Dougl.) in California and Oregon and those that parasitize south- western white pine (Pinus strobiformis Eng- elm.) in Arizona and New Mexico. Hawks- worth and Wiens (1965, 1970, 1972) described several new species of Arceutho- bium from Mexico and the Western United States and reported that many of Gill's host fonns could be distinguished morphologically and physiologically, including A. camp- ylopodum f. blumeri and f. cyanocarpum (A. Nelson) Gill. They separated f. blumeri into three species, A. californicum Hawksw. & Wiens, A. apachecum Hawksw. & Wiens, and A. blumeri A. Nelson, based on morphol- ogy, phenology, and geographic distribution. Arceuthobium apachecum exclusively para- 'School of Forestry, Northern Arizona University, Flagstaff, Arizona 86011. sitizes Pinus strobiformis and is distributed from the Santa Rita and Chiricahua Moun- tains of southern Arizona north to east cen- tral Arizona and west central New Mexico, with one population known from northern Coahuila, Mexico (Hawksworth and Wiens 1972, Mathiasen 1979). Arceuthobium blu- meri parasitizes Pinus strobiformis and Pinus ayacahuite var. brachyptera Shaw (Mexican white pine) and is distributed from the Hua- chuca Mountains, Arizona south through the Sierra Madre Occidental to southern Duran- go, Mexico, with one population known from Nuevo Leon, Mexico (Hawksworth and Wiens 1972, Mathiasen 1979). Arceuthobium apachecum and A. blumeri are morphologi- cally similar, but can be distinguished by shoot color, shoot height, growth habit, and number of perianth lobes of staminate flow- ers (Hawksworth and Wiens 1972). Few data, however, were available on the phenology of these species (Hawksworth and Wiens 1972), and because of their morphological similarity their specific status has been questioned (Kuijt 1973). This study was undertaken to provide more information on the morpho- logical and physiological characters of these two taxa. Materials and Methods Measurements and observations of mor- phological characters were made on 21 pop- ulations of A. apachecum and 18 populations of A. blumeri distributed throughout their ge- ographic ranges. Specimens examined were 120 March 1982 Mathiasen: Dwarf Mistletoe 121 collected by the author in 1975 and 1976 or were previously deposited at the U.S. Forest Service, Forest Pathology Herbarium, Fort Collins, Colorado (FPF) or the University of Arizona Herbarium, Tucson, Arizona (ARIZ). The specimens examined represent essen- tially all the material that has been collected for these species. Measurements and observa- tions were made for the following morpholo- gical characters: staminate and pistillate shoot height, pistillate shoot basal diameter, staminate and pistillate shoot color, and shoot growth habit (all measurements and observa- tions for dominant shoots of nonsystemic in- fections only); length and width of staminate spikes in summer; staminate flower diameter; perianth lobe number and color; mature fruit length, width, and color; mature seed length, width, and color. The original measurements of shoot height made for these taxa by Hawksworth and Wiens (1972) were avail- able and were included in obtaining mean, maximum, and minimum values for this character. Observations of flowering and seed dis- persal for both species were made in Arizona and New Mexico in the summer and fall of 1973 through 1976. The species, location, elevation, and date were recorded for each field observation, and flowering and seed dis- persal were each rated as not started, started but not near peak, near peak, past peak but not completed, and completed. Phenological data were examined by weekly periods for all observations from 1 July through 2 Novem- ber to determine the approximate periods of flowering and seed dispersal for both species. Results Mean heights of staminate and pistillate shoots of A. blumeri were approximately 1.5 cm greater than those of A. apachectim, and maximum shoot heights of A. bhimeri were also twice those of A. apachecum (Table 1). Nevertheless, analysis of shoot heights for these taxa using the method for comparing two sample means described by Cochran and Cox (1957) for samples with unpaired obser- vations and unequal variance indicates that the differences are not significant at the 5 percent level. Analysis of separate popu- lations from east central Arizona to southern Durango, Mexico, indicates that shoots are smallest for the northern populations of both species and largest for the southern popu- lations (Table 2). In addition, shoot heights are approximately the same for these taxa in southern Arizona (Table 2). Shoots of A. blu- meri vary in color from light green to straw or gray, and those of A. apachecum vary from yellow-green to blue or reddish. Both species parasitize Pinus strobiformis, but their growth habit is different on this host. Shoots of A. apachecum are consistently densely clustered around the host branch and may even completely obscure the branch. Shoots of A. bhimeri are more scattered and are rarely densely clustered on a branch. Lateral staminate spikes are larger for A. bhimeri (means 12 X 2 mm) than for A. apachecum (means 7x1 mm). The mean di- ameter of staminate flowers of A. bhimeri is larger than that of A. apachecum, but their Table 1. Comparison of selected morphological characters of Arceuthobium apachecwn and Arceuthobitirn blumeri. A. apachecum A. blumeri No. No. populations No. populations No. Character Mean Max. Min. sampled measured Mean Max. Min. sampled measured Shoot height (cm) Pistillate 4.7 9.5 2.5 21 254 6.1 18.0 3.0 18 218 Staminate 3.4 7.5 2.0 16 194 5.0 16.0 2.0 14 107 Shoot basal diameter (mm) (Pistillate) 1.7 4.4 0.8 21 234 2.0 3.4 1.2 18 193 Staminate flower diameter (mm) 2.9 4.2 2.2 10 296 3.2 4.4 2.0 6 146 Mature fniit (mm) Length 3.2 4.0 2.6 14 310 3.5 4.0 2.0 10 175 Width 1.9 2.4 1.4 14 310 2.0 2.4 1.6 10 175 122 Great Basin Naturalist Vol. 42, No. 1 size range is approximately the same (Table 1). Perianth lobe dimensions are similar also, but their color varies. In A. apachecum the perianth lobes are the same color as the male shoots, but in A. blumeri they are darker. These species also differ in the number of perianth lobes. Arceuthobium apachecum is predominantly 3-merous (65 percent), com- monly 4-merous (33 percent), and rarely 5- merous (2 percent); A. blumeri is pre- dominantly 4-merous (53 percent), less com- monly 3 or 5-merous (31 percent and 15 per- cent, respectively), and rarely 6-merous (1 percent). Comparison of perianth lobe num- ber in separate populations from east central Arizona to southern Durango, Mexico, in- dicates that 3-merous staminate flowers pre- dominate in the northern populations of A. apachecum and gradually change until 4- merous flowers predominate in most of the southern populations of this species (Table 3). Four-merous flowers predominate in the most northern populations of A. blumeri, but the few counts made for this species in Mexi- co indicate that 3-merous flowers may pre- dominate in the southern populations. Six- merous flowers were found only in the Ari- zona population of A. blumeri (Huachuca Mountains). Fruits are approximately the same size for both species (Table 1). Mean dimensions of mature seed were similar also (2.6 X 1.1 mm for A. blumeri and 2.3 X 1.0 mm for A. apachecum), but seeds of A. blumeri are dark green and those of A. apachecum are light green. Seeds from Mexican populations of A. blumeri were not available for examination, however. Table 2. Geographic variation in shoot height oi Arceuthobium apachecum and Arceuthobium blumeri. Location (Latitude) White Mountains, Arizona Mogollon Mountains, New Mexico (33°20'-34°10'N) Mangas Mountains, New Mexico (34'^5'N) Magdalena and San Mateo Mountains, New Mexico (34°N) Capitan Mountains, New Mexico (33°30'N) Pinaleno Mountains, Arizona (32°30'N) Santa Catalina Mountains, Arizona (32°25'N) Chiricahua Mountains, Arizona (31°50'N) Santa Rita Mountains, Arizona (31°40'N) Huachuca Mountains, Arizona (31°30'N) Sierra de Ajos, Sonora, Mexico (30°30'N) Chihuahua, Mexico (26°30'-29°30'N) Durango, Mexico (23°-26°N) Pistillate (cm) Mean Max. A. apachecum 4.0 5.0 Staminate (cm) Mean 3.1 No. _ populations Max. sampled 4.3 4.1 5.0 2.0 3.0 2 4.7 6.0 3,7 4.5 2 4.2 5.5 2.0 4.0 2 4.2 5.0 3.7 4.4 2 5.8 7.5 4.3 6.1 2 4.9 6.0 4.2 5.0 2 6.3 9.5 4.9 7.5 3 A. blumeri 6.4 8.5 4.2 8.0 3 6.0 7.5 - - 1 5.9 11.5 4.7 8.0 8 8.0 18.0 7.6 16.0 5 March 1982 Mathiasen: Dwarf Mistletoe 123 Arceutliobium blumeri consistently flowers earlier than A. apachecum (Fig. 1), but one of the most southern populations of A. apache- cum (Santa Rita Mountains, Arizona) flowers at approximately the same time as the most northern population of A. blumeri (Huachuca Moimtains, Arizona). These populations are separated by almost 40 miles. Arceutliobium apachecum disperses seed somewhat earlier than A. bhimeri (Fig. 1). Data on the flower- ing and seed dispersal periods of A. blumeri populations in Mexico are still inadequate, but tliey are evidently similar to the A. blu- meri population in Arizona (Hawksworth and Wiens 1972). Both species show altitudinal variation in flowering and seed dispersal. Lower populations flower earlier than higher populations in the same mountain ranges in southern Arizona and the reverse is true for seed dispersal. Flowering of A. apachecum begins as early as mid-August in the northern populations of this species, but does not start imtil early September in the southern popu- lations observed. Annual climatic variations also influence the phenology of these species. Seed dispersal starts later in years with a late fall. Discussion Arceuthobium apachecum and A. blumeri are morphologically similar, but differ in col- or of staminate and pistillate shoots, growth habit, dimensions of lateral staminate spikes, color of perianth lobes, and seed color. Shoot height and number of perianth lobes appear to be discontinuous characters also, but anal- ysis of different populations indicates that considerable geographic variation occurs in these characters. Although mean shoot heights are different for these species, shoots are shortest in the northern populations of A. apachecum, tallest in the southern popu- lations of A. blumeri, and approximately the same near the geographic boundary between these taxa in southern Arizona. Variation in the number of perianth lobes appears to fol- low a geographic pattern also, but more in- formation is needed for Mexican populations of A. blumeri. Six-merous flowers are only known for A. blumeri, however. Periods of flowering and seed dispersal for A. apachecum and A. blumeri are slightly dif- ferent, although one population of A. apache- cum does flower at approximately the same Table 3. Geographic variation in number of perianth lobes of Arceuthobium apachecum and Arceuthobium bhimeri. Location (Latitude) 3 4 5 6 No. populations sampled No. measured A. apachecum (%) White Mountains, Arizona (33°20'-34°10'N) 81 19 0 0 6 390 Pinaleno Mountains, Arizona (32°30'N) 68 30 2 0 2 50 Chiricahua Mountains, Arizona (31°50'N) 42 54 4 0 2 50 Santa Catalina Mountains, Arizona (32°25'N) 54 42 4 0 2 480 Santa Rita Moimtains, Arizona (31°40'N) 43 48 8 0 3 500 A. bhimeri (%) Huachuca Mountains, Arizona (31°30'N) 30 53 16 1 3 700 Chihuahua, Mexico (26°30'-29°30'N) 40 58 2 0 5 50 Durango, Mexico (23°-26°N) 55 44 1 0 5 100 124 Great Basin Naturalist Vol. 42, No. 1 time as A. blumeri in Arizona. Latitudinal variation, however, may influence these dif- ferences in flowering and seed dispersal. Ar- ceuthohium blumeri has a more southerly range and flowers before A. apachecum, but disperses seed slightly later. This pattern is evident in the elevational variation observed for both taxa where lower populations begin flowering before higher populations in the same mountain ranges, but disperse seed sooner at the higher elevations. Scharpf (1965) reported a similar altitudinal relation- ship for flowering and seed dispersal of A. abietinum Engelm. ex Munz in California. Northern populations of A. apachecum start seed dispersal before southern populations of this species in Arizona, which also suggests that latitude influences this character. There fore, the slight differences in the flowering and seed dispersal periods of these species, and possibly other fall-flowering species in the Series Campyhpoda Hawksw. & Wiens (Hawksworth and Wiens 1970, 1972) may be a result of climatic fluctuations, possibly tem- perature variation (Scharpf 1965), associated with latitudinal, altitudinal, or seasonal dif- ferences. Variation in the flowering and seed dispersal periods of dwarf mistletoes is com- plex and deserves more study before phenol- ogy can play an important role in the classifi- cation of fall-flowering species of Arceuthobium. Environmental factors may influence other physiological characters of dwarf mistletoes. The consistent inducement of witches' brooms by A. apachecum and the rare in- ducement of brooms by A. blumeri was con- sidered as a taxonomically significant dis- continuity between these species by Hawksworth and Wiens (1972). Some popu- lations of both species rarely cause brooms at lower elevations, however, but frequently cause brooms at higher elevations (Mathiasen 1979). The reason for this apparent relation- ship between witches' broom formation and elevation is unknown, but the rare in- ducement of brooms by A. blumeri reported by Hawksworth and Wiens is not typical of that species. Geographic variation in morphological and physiological characters of dwarf mistletoes may be influenced by variation in their host population as well as parasitism of different hosts. Shoot height of dwarf mistletoes is di- rectly related to variation in host vigor (Hawksworth 1960, Hawksworth and Wiens A. BLUMERI A. APACHECUM A. BLUMERI A. APACHECUM ANTHESIS ( 26) 1(84) SEED DISPERSAL j (26) (88) 15 31 15 31 15 30 15 31 JULY AUGUST SEPTEMBER OCTOBER Fig. 1. Approximate periods of anthesis and seed dispersal of Arceuthobium blumeri and Arceuthobium apache- cum. Peak periods are shown by solid bars. Number of observations are in parentheses. March 1982 Mathiasen: Dwarf Mistletoe 125 1972) and may itself influence other charac- ters of a dwarf mistletoe. The southern popu- lations of A. blumeri parasitize Finns ayaca- huite, which is one of the largest pines in Mexico (Loock 1950, Mathiasen 1979). Pimis ayacahuite commonly occurs in moist local- ities, with deep, well-drained soils that are conducive to maintaining a vigorous growth rate (Loock 1950). Therefore, the large shoot heights found in the southern populations of A. bhimeri may be related to the parasitism of a more vigorous host population. The tax- onomic relationships between P. ayacahuite and P. strobifonnis are uncertain. Morpho- logical and physiological variation has been reported in different populations of P. strobi- fonnis (Andresen and Steinhoff 1971) and the various taxonomic treatments of this species in different parts of its range suggest that it is quite variable (Steinhoff and Andresen 1971). Critchfield and Little (1966) considered all white pine populations from Durango, Mexi- co, northward to the southwestern United States to be P. strobifonnis. Nevertheless, other investigators believe that many of the white pine populations in northern Mexico are typical of P. ayacahuite (Martinez 1948, Loock 1950, Mathiasen 1979). Further tax- onomic studies of the white pine populations represented in the southwestern United States and northern Mexico are needed and may provide additional information con- cerning the relationships between A. blumeri and A. apachecum. Hawksworth and Wiens (1972) originally considered that the dwarf mistletoes para- sitizing P. strobiformis might represent a single variable taxon. They finally concluded that A. apachecum and A. blumeri warranted separate taxonomic status at the specific lev- el, because they believed the morphological and physiological differences they detected between these taxa were geographically con- sistent. Hawksworth and Wiens (1972) con- sidered two criteria as the most important factors in delimiting species of Arceutho- bium: (1) species maintain their morphologi- cal integrity when parasitizing species other than their principal hosts, and (2) species of Arceuthobium are often sympatric but do not show evidence of hybridization. It is not known if A. apachecum and A. blumeri meet these criteria because they are not sympatric and do not parasitize any species other than their principal hosts (Hawksworth and Wiens 1972, Mathiasen 1979). Hawksworth and Wiens (1972), however, reported that these taxa maintain their morphological integrity when grown under common greenhouse con- ditions. In addition, chemical analysis of the shoots of these species has shown there are consistent differences in their phenolic chem- istry (Hawksworth and Wiens 1972, Craw- ford and Hawksworth 1979). Artificial hy- bridization of A. apachecum and A. blumeri has been attempted, but the results were in- conclusive (Mathiasen, unpubl. ms.). Crosses between A. apachecum (staminate) and A. blumeri (pistillate) were destroyed by a wild- fire in the Huachuca Mountains, Arizona, in 1977 before results could be assessed. Crosses between A. blumeri (staminate) and A. apachecum (pistillate) in the Santa Catalina Mountains, Arizona, resulted in no successful fruit set, but the control crosses (A. apache- cum [staminate] X A. apachecum [pistillate]) yielded very poor fruit set. These in- vestigations must be repeated before their re- sults can be considered as evidence these spe- cies are reproductively incompatible. Although geographic variation occurs in some of the morphological and physiological characters used by Hawksworth and Wiens to delimit A. apachecum and A. blumeri, the results of this study indicate there are several geographically consistent morphological dis- continuities between these taxa. I believe these morphological differences are tax- onomically significant and suggest that these dwarf mistletoes should be given separate taxonomic recognition. Nevertheless, these populations may represent intermediate stages of gradual evolutionary divergence, and perhaps their recognition at the-^ syb- specific level would be more representative of their natural relationship. Hawksworth and Wiens (1972) defined subspecies of Arceuthobium as "geographi- cally restricted populations, delimited by rel- atively few but consistent variations." The dwarf mistletoe populations parasitizing Pinus strobifonnis appear to more closely conform to these criteria than those used by Hawksworth and Wiens to define species of Arceuthobium. Examination of the dis- continuities separating currently recognized 126 Great Basin Naturalist Vol. 42, No. 1 subspecies of Arceuthobium (Hawksworth and Wiens 1970, 1972, 1977) indicates there are more morphological and physiological differences between these taxa than between A. apachecum and A. hlunieri. Several spe- cies in the Series Campylopoda, however, in- cluding A. apachecum and A. blumeri, are delimited by relatively few morphological and physiological differences, so tiie classifi cation of taxa in this series is relatively con- sistent. Because species and subspecies of Ar- ceuthobium differ only in the number of dis- continuities between them (Hawksworth and Wiens 1972), the elevation of currently rec- ognized subspecies to specific rank or the separation of some species into subspecies would alleviate the apparent inconsistencies I feel exist in the present classification of Ar- ceuthobium. Consistency in the classification of a group is desirable and changes in the rank of taxa for motives of consistency or for achieving a more balanced natural classifica- tion are justified (Davis and Heywood 1963). Isozyme analyses and additional field studies of dwarf mistletoe populations in Mexico and the western United States are in progress, and the results may provide evidence that changes in the rank of some taxa would cre- ate, a more natural and consistent classifica- tion of the genus (F. G. Hawksworth and Dan Nikrent, pers. comm., 1981). Therefore, I do not feel that a change in the rank of A. apachecum or A. blumeri should be consid- ered until additional information is available concerning their natural relationships and their relationship with other species in the Series Campylopoda. Stability in the classification of such an economically important group as Arceutho- bium is desirable; yet it is doubtful that this can be achieved in the near future because little is known about the ranges and natural relationships of the recently discovered Mexi- can and Central American species (Hawks- worth and Wiens 1972, 1977, 1980). More than one-half of the taxa now recognized have been described in the last 10 to 15 years, so much critical work on the entire genus will be required before a more stable classification of Arceuthobium can be achieved. Acknowledgments This research was supported by the U.S. Forest Service, Rocky Mountain Forest and Range Experiment Station, and the Univer- sity of Arizona, Department of Plant Pathol- ogy. I would particularly like to thank F. G. Hawksworth for his support and constructive comments regarding the final manuscript. Our discussions concerning the taxonomy of Arceuthobium have greatly contributed to this work. Literature Cited Andresen, J. M., AND R. J. Steinhoff. 197L The tax- onomy of Pinits flexilis and Pinus strobiformis. Phytologia 22: 57-70. Cochran, W. G., and G. M. Cox. 1957. Experimental designs, 2d ed., John Wiley & Sons, Inc., New York. Crawford, D., and F. G. Hawksworth. 1979. Flavo- noid chemistry of Arceuthobium (Viscaceae). Brittonia 31:212-216. Critchfield, W. B., and E. L. Little. 1966. Geo- graphic distribution of the pines of the world. U.S. Dept. Agric, Misc. Publ. 991, 97 pp. Davis, P. H., and V. H. Heywood. 1963. Principles of angiosperm taxonomy. Oliver and Boyd, Prince- ton, New Jersey. Gill, L. S. 1935. Arceuthobium in the United States. Connecticut Acad. Arts and Sci. Trans. 32: 111-245. Gray, A. 1850. Plantae Lindheimerianae. Part II. Boston J. Natur. Hist. 6: 141-240. Hawksworth, F. G. I960. Growth rate of dwarf mis- tletoe infections in relation to the crown class of the host U.S. Dept. Agric. For. Serv., Res. Note RM-41, 4 p.. Rocky Mtn. For. and Range Exp. Stn., Fort Collins, Colorado. Hawksworth, F. G., and D. Wiens. 1965. Arceutho- bium in Mexico. Brittonia 17: 213-238. 1970. New taxa and nomenclatural changes in Arceuthobium (Viscaceae). Brittonia 22: 265-269. 1972. Biology and classification of dwarf mis- tletoes (Arceuthobium). U.S. Dept. Agric. For. Serv., Agric. Handb. 401, 234 pp. 1977. Arceuthobium (Viscaceae) in Mexico and Guatemala: additions and range extensions. Brit- tonia 29: 411-418. 1980. A new species of Arceuthobium (Viscaceae) from central Mexico. Brittonia 32: 348-352. KuijT, J. 1973. Review of "Biology and classification of dwarf mistletoes (Arceuthobium)" by F. G. Hawksworth and D. Wiens. 1972. Madrono 22: 34-35. LoocK, E. E. M. 1950. The pines of Mexico and British Honduras. Union South Africa Dept. For. Bull. 35, 244 pp. Martinez, M. 1948. Los pinos Mexicanos. Ed. 2 Mexico, 361 pp. March 1982 Mathiasen: Dwarf Mistletoe 127 Mathiasen, R. L. 1979. Distribution and effect of dwarf mistletoes parasitizing Pinus stwhiformis in Ari- zona, New Mexico, and northern Mexico. South- western Natur. 24: 455-461. ScHARPF, R. F. 1965. Flowering and seed dispersal of dwarf mistletoe [Arceuthobium camptjlopodum) in California. U.S. Dept. Agric. For. Serv., Res. Note PSW-68, 6 p., Pac. Southwest For. and Range Exp. Stn., Berkeley, California. Steinhoff, R. J., AND J. W. Andresen. 1971. Geographic variation in Pinits flexilis and Pinus stwhiformis and its bearing on their taxonomic status. Silvae Genetica 20: 159-167. Watson, S. 1880. Botany in California. II. Pages 105-107 in Apetalea, Gymnospermae, Mono- cotyiedonous, or endogenous plants, Crypto- gamous plants. Welsh, Bigelow, and Co., Cam- bridge, Massachusetts. 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. 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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: Rosaceae. Stanley L. Welsh 1 Seasonal foods of coyotes in southeastern Idaho: a multivariate analysis. James G. MacCracken and Richard M. Hansen 45 Stnictiire of Alpine plant communities near King's Peak, Uinta Mountains, Utah. George M. Briggs and James \. MacMahon 50 Observations on the reproduction and embryology of the Lahontan tui chub, Gila bicolor, in Walker Lake, Nevada. James J. Cooper 60 The prevalence of Echinococcus granulosus and other taeniid cestodes in sheep dogs of central Utah. Lauritz A. Jensen, Ferron L. Andersen, and Peter M. Schantz 65 Growth of juvenile American lobsters in semiopen and closed culture systems using formulated diets. S. R. Wadley, R. A. Heckmann, R. C. Infanger, and R. W. Mickelsen 67 Diameter-weight relationships for juniper from wet and dry sites. T. Weaver and R. Lund 73 A description of Timpie Springs, Utah, with a preliminary survey of the aquatic macrobiota. Thomas M. Baugh, Michael A. Nelson, and Floyd Simpson 77 Vegetation and soil factors in relation to slope position on foothill knolls in the Uinta Basin of Utah. Miles O. Moretti and Jack D. Brotherson 81 Weather conditions in early summer and their effects on September blue grouse (Dendragapus obscurus) harvest. Joy D. Cedarleaf, S. Dick Worthen, and Jack D. Brotherson 91 Description of the female of Phalacropsylla hamata (Siphonaptera: Hys- trichopsyllidae). R. B. Eads and G. O. Maupin 96 First record of pygmy rabbits (Brachylagus idahoensis) in Wyoming. Thomas M. Campbell III, Tim W. Clark, and Craig R. Groves '. 100 Paspalum distichum L. var. indutum Shinners (Poaceae). Kelly Wayne Allred 101 Local floras of the southwest, 1920-1980: an annotated bibliography. Janice E. Bowers 105 The relation between species numbers and island characteristics for habitat islands in a volcanic landscape. Steven H. Carter-Lovejoy 11.3 Taxonomic studies of dwarf mistletoes {Arceuthobium spp.) parasitizing Pinus strobi- formis. Robert L. Mathiasen 120 THE GREAT BASIN NATURALIST Volume 42 No. 2 June 30, 1982 Brigham Young University 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 Museimi, 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. 7-82 650 60479 ISSN 017-3614 MU! H'^T The Great Basiii Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 42 June 30, 1982 No. 2 UTAH PLANT TYPES-HISTORICAL PERSPECTIVE 1840 TO 1981- ANNOTATED LIST, AND BIBLIOGRAPHY St;uiley L. Welsh' .\bstract.— Reviewed are the 144 collectors and 167 authors of 1073 Utali vascular plant types for the period from 1840 to 1981. Historical perspective yields evidence of geography of collection activities by botanists who pene- trated the boundaries of the state, and of shifting centers of emphasis in the study of Utah plants from a classical tax- onomic standpoint. Philosophy and influence of contemporary authors and collectors is discussed as they affect tax- onomv of Utah plants in our first 142 years. A short biographical account of Marcus Eugene Jones and his interaction with other American botanists is included. The annotated checklist includes bibliographical citations, type locality data, collector, and place of deposition. Acknowledgments It is impossible to complete a work as complex and tedious as this without the help of otiiers. I thank the curators of the herbaria visited by me for their hospitality and will- ingness to help in this work. They are as fol- lows: Elizabeth McClintock and John Thomas Howell (CAS); Robert Thorne (RSA); Ronald Hartman (RM); Patricia K. Holmgren and Rupert C. Bameby (NY); George Russell (US); Lois Amow and Beverly Albee (UT); Mary Barkworth (UTC); Theodore J. Cro- vello (NDG); and Richard Pohl and Duane Isely (ISC). I am indebted to George Knaphus and Lois Tiffany who made it possible for me to visit the site of Jones Grove, where Marcus E. Jones lived with his family near Grinnell, Iowa. Acknowledged also is the help of Mrs. Ralph Fleener, who resides currently on the former Jones's property, for the checking of records and dates on markers in the cemetery at Grinnell. Kaye Thorne, my colleague at the herba- rium of Brigham Young University, gave en- couragement and help at all times. Mark L. Gabel aided me with research on the life of Charles Christopher Parry. To him I am grateful. To Mr. Charles Walker, archivist for the family of Publius Virgilius and Lavinia (Bur- ton) Jones, I am grateful for the access he provided me to a published family history in- cluding accounts of Marcus E. Jones and his wife, Anna Elizabeth Richardson. Also, he provided me original letters written by both of them, and by a nephew of Jones, Arthur J. Jones, who accompanied him on attenuated field trips by wagon. These have given me in- sight into Jones's family life and have allowed me to present a portion of the life-style of his wife and family during his intensely goal- oriented life. I shall be forever indebted to Mr. Walker and his family for this insight. 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 129 130 Great Basin Naturalist Vol. 42, No. 2 Introduction All taxonomists, and others who work with plant names, have been hampered by lack of a summary treatment of Utah plant types, their collectors, authors, and places of deposi- tion. The specimens serve to typify the names applied to them by the authors who named and described them. The types are the name-bearing specimens and are in- dispensible in interpretation of taxonomic hmits and concepts; they are necessary for nomenclatural considerations. There are several kinds of types, but only holotypes and their duplicates, the isotypes, have been included uniformly in the follow- ing treatment. I have not tried to distinguish holotypes from isotypes in the designations of place of deposition; that is a taxonomic con- sideration which is left to the monographer. In many instances the name bearing speci- mens were not designated as types by the au- thors and I have followed tradition in citing the original materials as types. Many of the older names have been subjected to lecto- typification. Where this has been obvious, mainly in taxonomic revisions or as indicated on some herbarium annotations, I have fol- lowed that usage. Lectotypification, how- ever, is subject to interpretation, and the specimens cited might ultimately be excluded from consideration as lectotype or isolecto- type. Too, some materials have been cited as cotypes or syntypes. When a portion of the original collection cited is from Utah, I have included that material as a type. It might have been designated at a lower level, i.e., as paratype, by some author or authors un- known to me. Designation of types is a function of rules established mainly in the 20th-century, and early authors were not impressed with the necessity of typification. Mention of speci- mens to vouch for newly described taxa was often only incidental. Most of the types serv- ing as bases for taxa described in the 19th- century are now interpreted through lecto- typification. Purists often are involved in polemical stRiggles over selection of the cor- rect portion of the specimens cited (or not cited) as the lectotype. I have not searched all the literature in which lectotypes have been designated; that is the role of the monographer. I have tried to avoid inclusion of obvious paratypes. These are specimens cited with the original description, but they are not du- plicates of the holotype. Mostly paratypes do, indeed, represent the same taxon as the holotype or lectotype. They are the next most important elements in typification; but sometimes they represent extraneous mate- rials, and might belong to quite different taxa. The main reason for not citing them here involves the endless listing of specimens. They are very numerous, and will be most important to those undertaking revisions and monographs. Information has been gathered from exam- ination of type collections at the U.S. Nation- al Museum (US, 500 + ); Rocky Mountain Herbarium (RM, 150 + ); California Academy of Science (CAS, 120+ ); New York Botanical Garden (NY, 550 + ); Rancho Santa Ana (POM, 240 + ); Iowa State University (ISC, ca 100); University of Utah (UT, 100+ ); Utah State University (UTC, 150 + ) and Brigham Young University (BRY, 250 + ). Only se- lected material has been examined from Gray Herbarium (GH), Carnegie Museum (CM), University of California, Berkeley (UC), Notre Dame University (NDG), and Missouri Botanical Garden (MO). Information on types at the Greene Herbarium at Notre Dame University was taken from a computer print- out provided through the generosity of Pro- fessor Theodore J. Crovello. Literature has been reviewed for most of the citations, but some have been unavailable. Other citations still await procedures entailed through inter- library loans. Type locality information has been abbre- viated in some instances to conform to the computer format. Localities of deposition, or supposed deposition, are indicated by the standard abbreviations. Where I have seen the type, the abbreviation is followed by an exclamation mark (e.g., US!). Where the ab- breviation stands alone, the place of deposi- tion is taken from literature, or from infer- ence with other collections or names published by the same author, but the speci- men has not been seen by me. Although portions of more than two dec- ades have passed since this work was begun, the data are still incomplete for many of the plant names. I apologize for that lack of June 1982 Welsh: Utah Plant Types 131 completeness, and for the many errors that will become apparent as one uses the list. Some of the problems are apparently in- soluble. Plant collectors and authors seldom have been instilled with sufficient foresight to supply all the information necessary for adequate intrepretation of specimens, local- ities, dates, and other data. Brevity was con- sidered to be a virtue by the collectors (along widi an almost impossible, if unstated, belief that others would know what they knew), even when that brevity left out such informa- tion as state, locality within the state (or ter- ritory), political subdivision of the state, and any other data. Herbarium labels are the sources of infonnation provided by plant col- lectors, and they should have contained cer- tain basic data. Some of the deficiency, such as lack of place names and surveys, is not the fault of the early workers. Boundaries of the state or territory of Utah were unknown until long after the final reduction of the original Territory of Deseret to its present size in 1868. When one went south from St. George, he was never quite sure when he had entered into Arizona. There is little excuse, however, for the Palmer and Parry labels with only "Southern Utah, Northern Arizona, &c." or "Southern Utah, &c." as the only locality data. In some instances the localities were known only to the collectors, whose passing has denied us of the possibility of ever knowing. Even when data were provided on labels, the author who subsequently published the name of the plant often failed to cite it. Many of the early publications, and some of the later ones, convey little in the way of ac- ciu^ate data on collector, number, date, local- ity, or place of deposition of the specimens. Some specimens were taken by obscure col- lectors, and the locality of collection, dates, and other information remains in doubt. Localities and dates for some of the plant names have been interpolated by reference with other collections. This is especially the case with county designations; boundaries have changed and new counties have been created since the earliest collections were made, and early collectors were not sure which county they were working in when some collections were made. Some pioneer towns and villages have passed into history. and others have undergone name changes. Present knowledge of distribution allows for accurate placement, at least to county level, of some materials which were provided with vague or misleading locality information. Problems associated with collections of certain individuals will be discussed in the historical review presented below. It is believed that the shortfalls in data do not obscure the basic picture of plant explor- ation in Utah or of the story of publication of taxa based on Utah types. This attempt at analysis of type data in no way detracts from the larger scene of plant studies which re- sulted from the naming of plants based on types from other areas. The flora of Utah is replete with names of plants with circum- boreal and other extra Utah representation dating to 18th- and early 19th-century collec- tors and authors. Acknowledged also are the contributions to botany of workers whose main thrust did not include classical taxonomy. Numerous botanists have contributed in ecology, phys- iology, anatomy, morphology, genetics, and other sophisticated fields of endeavor. Each of those areas has allowed for a greater un- derstanding of the taxonomy of the state. Some investigators contributed to more than one of the major botanical disciplines, or to taxonomy of nonvascular plants, not treated herein. Their exclusion from the lists of col- lectors and authors, or their low placement in the lists does not imply that the contributions are not great. The measure herein is based on contributions in the strictly classical sense. Neither should naming and description of plants be expected to continue indefinitely. But, the botany of the state is not yet eluci- dated, and much further work is indicated. This work should not be thought of as repre- senting the final word; certainly many cita- tions of Utah types have been overlooked. Collectors of Utah Types The number of vascular plant types report- ed for Utah between 1840 and 1981 is ap- proximately 1073. Exact numbers are impos- sible due to such equivocal label data as "Southern Utah, Northern Arizona, &c." for some collections. The total number that might be in error is thought to be small, and 132 Great Basin Naturalist Vol. 42, No. 2 for the purposes of this paper they seem not to affect the overall picture. The types have been taken by 144 collectors acting singly or in combination with one or more collabora- tors (Table 1). Possibly other collectors are obscured by citations where collection data are lacking. Authors include some 167 indi- viduals acting singly or in combination, and will be discussed later. A summary of collections of Utah plant types by decade is presented in Table 2. Earliest types taken from Utah were ap- parently those by John Charles Fremont in 1843 [Perchiton occidentale Torr. & Frem. = Atriplex canescens (Pursh) Nutt.] on his first expedition in the American West, wherein he explored the vicinity of the Great Salt Lake. The types of Gilia stenothijrsa T. & G. and Senecio multilobatus T. & G. were collected by Fremont apparently during 1845 in the Uinta Basin. The 1850s were marked by the efforts of Captain Howard Stansbury, F. Creutzfeldt, Henry Engelmann, and Edward Griffin Beckwith. Stansbury explored the Great Salt Lake in 1850; Creutzfeldt and Beckwith were associated with the ill-fated Gunnison expedition of 1853-54. Creutzfeldt was the botanist for the expedition, whose duty was to survey a route for a railroad to the Pacific (Beckwith 1854), and he was killed along with John Williams Gminison and most of the others of the party in an am- bush by Indians near Delta on 26 October 1853. Beckwith, who was leading another portion of the expedition at the time of the disaster, was named to succeed Gunnison. Creutzfeldt had collected types of two Table 1. Collectors of Utah plant types. Table 1 continued. Collectors Number Years of types active Collectors Number of types Years active Al-Shehbaz, I.A. (b.?) 1 1969 Atwood, N.D. (b. 19.38) 3 1970-76 Atwood, N.D. (b.l938) & L.C. Higgins (b.l936) 1969-73 Atwood, N.D. (b.l938) et al. 1971 Bailey, V.O. (1864-1982) 1891 Baker, J.G. (1834-1920) 189,5 Baker, M.S. (1868-1961) 19.36 Ball, C.R. (1873-19.58) 1908 Barklev, F.A. (b.?) & M.J. Reed (b.?) 1 1939 Barneby, B.C. (b.l911) Beck, D E. (1906-1967) Beckwith, E.G. (1818-1881) Benson, L.D. (b.l909) Bishop, P.M. (184.3-19.33) Brandegee, T.S. (1843-1925) Burke, M.(b.?) Carlton, E.C. (b.?) & A.O. Garrett (1870-1948) Carrington, J. (b.?) Carter Clawson Clover, E.U. (b.l897) Clover, E.U. (b. 1897) ik M.L. Jotter (b. 1914) Clute, W.N. (b.l869) Cottam, W.P. (b.l894) Coville, F.V. (1867-1937) & I.T. Tidestrom (1864-1956) Creutzfeldt, F. (d. 18,53) Cronquist, A.J. (b.l919) Despain, K. (b.l947) Eastwood, A. (1859-1953) Eastwood, A. (1859-19,53) & J.T. Howell (b. 190.3) Eaton, D.C. (1834-189,5) Eggleston, W.W. (186.3-1935) Ehrendorfer, F. (b.l927) & H.C. Stutz(b.l918) Engelmann, H. (1831-1899) Fallas Ferguson & A.M. Ottley (b.l882) Flowers, S. (1900-1968) Fremont, J.C. (181,3-1890) Garrett, A.O. (1870-1948) Gentry, J.L. (b.?) Gilbert, G.K. (1843-1916) Goodding, L.N. (b.l880) Goodman, G.J. (b.l904) &C.L. Hitchcock (b. 1902) Goodrich, S. (b.l943) Gould, F.W. (b. 191.3) Graham, E.H. (b.l902) Hamilton, J.W. (b.?) & O.A. Beath(b.l884) Hanson, C.A. (b.l935) Harkness, S.J. (b.?) Harrison, B.F. (b.l908) Heil, K.D. (b.l941) Hermann, F.J. (b.l906) Hermann, A. (b.?) Higgins, L.C. (b.l936) Hitchcock, C.L. (b.l902) Holmgren, A.H. (b.l912) et al. Holmgren, A.H. (b.l912) & S.S. Tillett (b.?) Holmgren, N.H. (b.l937) et al. 5 1956-66 1 19,38 4 1854 4 1948-49 4 1872-73 2 1873-75 1 1932 3 1905 2 1857 1 1938 2 1935 1 1940 1 1940 1 1919 8 1929-37 1 1908 2 1853 4 1961 1 1978 ,361892-1941 9 1933-41 3 1869 1 1912 1 1959 1 1859 1 1928 1 p 1 1933 4 1843-45 251894-1940 1 1968 1 1901 21 1902 3 1930 2 1980 1 1941 6 1935-37 1 1952 3 1960-61 1 1902 2 1939-50 2 p 3 1933 1 1953 3 1968 1 1949 4 1947-74 1 1953 10 1965-79 June 1982 Table 1 continued. Welsh: Utah Plant Types Table 1 continued. 133 Collectors Holmgren, N.H. (b.l937) &P.K. Holmgren (b. 1940) Holmgren, N.H. (b. 1937) &J.L. Reveal (b. 1941) Huffman Johnson, J.E. (1817-1882) Jones, M.E. (1852-1934) Kearney, T.H. (1874-1956) & H.L. Shantz (1867-1958) Klein, W. (b.?) Kuntze, C.E.O. (1843-1907) Larson Leonard, F.E. (1866-1922) Linford, J.H. (b.l863) Maguire, B. (b.l904) Maguire, B. (b.l904) & H.L. Blood (b.?) Maguire, B. (b.l904) & A.H. Holmgren (b.l912) Maguire, B. (b.l904) & R. Maguire (b.?) Maguire, B^ (b.l904) et al. Marsh Mathias, M.E. (b.l906) McKelvey, S.D. (b.l883) Mulford,'A.L (b.?) Neese, E. (b.l934) & S. Peterson (b.?) Neese, E.(b. 1934) & S.L. Welsh (b.l928) Neese, E.(b. 1934) &S. White (b. 1955) Nelson, A. (1859-1952) Newberry, J.S. (1822-1892) Nord & Sargent Osterhout, G.E. (1858-1937) Ostler, W.K.(b. 1951) &D.C. Anderson (b. 1948) Ownbey, G.B. (b.l916) Palmer^ E. (1831-1911) Pammel, L.H. (1862-1931) & R.E. Blackwood (b,?) Parry, C.C. (1823-1890) Payson, E.B. (1893-1927) Peabodv, F.J. (b.l948) Pennell', F.W. (1886-1952) Pilsbry Plummer, P. (b.l911) Porter, C.L. (b.l889) Purpus,C. A. (1851-1941) Reading Reveal,"j.L. (b.l941) Reveal, J.L.(b. 1941) & G. Davidse (b.?) Reveal, J.L. (b.l941) & N.H. Holmgren (b.l937) Reveal, J.L. (b. 1941) & C. Reveal (b.?) Number of types Years active 4 1970-78 7 1965-69 3 1953 2 1870 3051879-1923 2 1912 1 1959 2 1874 1 1934 11 1883-85 9 1897 26 1932-47 1937 1946 2 19.32 4 19.36-42 3 1945 2 1929 2 1934 3 1898 1978 1979 1 1977 4 1922 2 1959 1 1927 1 1925 1 1978 1 1954 58 1870-77 2 1902 38 1874 5 1914-26 1 1976 1 1915 1 1925 1 1939 3 1939-50 4 1897-99 1 ■p 3 1964-74 1967 1964 3 1967-75 Number Years Collectors of types active Reveal, J.L. (b. 1941) & S.L. Welsh (b. 1928) 1 1966 Ricker, P.L (1878-1973) 1 1917 Ripley, H.D.D. (b.?) &R.C. Barneby(b.l911) 11 1942-47 Rollins, R.C.(b.l911) / 1937-79 Rollins, R.C. (b.l911) & T.S. Chambers (b.?) 1 19.38 Rydberg, P.A. (1860-1931) 81895-1911 Rydberg, P.A. (1860-1931) & E.G. Carlton (b.?) 22 1905 Rydberg, P.A. (1860-1931) '& A.O.Garrett (1870-1948) 24 1911-13 Sampson, A.W.(b. 1884) 1 p Shands, W.A. 1 1931 Shultz, L. (b.l946) & J. Shultz (b.?) 1 1980 Siler,A.L.( 1824-1898) 6 1873-75 Smith, C. P. (1877-1955) 6 1909-25 Stansbury, H. (1806-1853) 11 1850 Stanton, W.D.(b. 1900) 2 1930-32 Stokes, S.G. (1868-19.54) 4 1900-34 Stoutamire, W.P. (b.?) 1 1957 Thompson, E.L.P. (b.?d.?) 15 1872 Tidestrom, I. (1864-1957) 13 1901-19 Tompkins, P.W. (b.?) 1 1939 Tracy, S.M. (1845-1922) 3 1887-91 Tracy, S.M. (1845-1922) & Evans 1 1887 Trelease, W. (1857-1945) 1 1889 Vicker 1 1901 Walker, E.B. (b.?) 5 1912 Ward, L.F. (1841-1913) 30 1875 Warner, L. (b.?) 1 1957 Watson, S. (1826-1892) 100 1869 Watson, T.J. (b.?) 1 1971 Welsh, S.L. (b.l928) 12 1966-80 Welsh, S.L. (b.l928) & N.D. Atwood(b.l938) 3 1970-72 Welsh, S.L. (b.l928) &G. Moore (b. 1917) 2 1963-76 Welsh, S.L. (b.l928) & J.R. Murdock(b.l921) 1 1975 Welsh, S.L. (b.l928) & K.N. Taylor (b. 1955) 2 1977 Welsh, S.L. (b.l928) & B.J. Welsh (b. 1961) 1 1977 Welsh, S.L. (b.l928) &S.L.T. Welsh (b. 1930) 5 1969-75 Welsh, S.L. (b. 1928) etal. 5 1969-75 Wetherill, B.A. (1861-1950) 3 1895-97 Wheeler, G.M. (b.l842) 3 1872 Williams, L.O. (b.l908) 1 1932 Winkler 1 ? Woodbury, L.A. (b.?) 1 1977 TOTAL 1073 134 Great Basin Naturalist Vol. 42, No. 2 Eriogonum taxa in the Green River vicinity. Beckwith took the type of a handsome Astra- galus, which bears his name. Several taxa were later dedicated to the memory of Gun- nison, but unfortunately none of our Utah materials bears the name of Creutzfeldt. Eng- elmann was meteorologist and botanist with the Simpson expedition (Ewan 1950) and took the type of Echinocactus whipplei var. spinosior Englemann (named by his brother George Engelmann) somewhere west of Camp Floyd in August 1859. The first woman collector of Utah types appeared on the scene in the 1850s. Jane Carrington (a Mormon lady) collected some 59 species in the basin of the Great Salt Lake in 1857 (Reveal 1972). She was apparently the first woman to collect plants in the terri- tory since the arrival of the Mormon pio- neers, a decade earlier. Two of her collec- tions are the basis of types. They were named by Elias Durand (1860) as Erysimum asperum var. purshii and Acerates decumhens. Perhaps this person is Jane Maria Carrington (26 Feb- ruary 1840, Wiota, Wisconsin), daughter of Albert Carrington and Rhoda Maria Woods, who married Brigham Young, Jr. in March of 1857. Albert Carrington was prominent in early day affairs of the territory of Deseret, serving as editor of the Deseret News (Cool- ey 1980). Albert Carrington accompanied Stansbury on his exploration of the Great Salt Table 2. Decade of collection of Utah types, number of collectors,and average number of types per collector. Number of Number of Decade Types Collectors T/C 1840-49 4 2 2.0 1850-59 23 5 4.4 1860-69 103 2 51.5 1870-79 184 10 18.4 1880-89 54 5 10.6 1890-99 259 9 28.8 1900-09 105 13 8.0 1910-19 60 13 4.5 1920-29 19 9 2.1 1930-39 85 31 2.7 1940-49 46 11 4.1 1950-59 12 11 1.1 1960-69 47 16 2.9 1970-79 52 24 2.2 1980-81 6 2 3.0 Without date 20 4 _ Total 1073 144 7.4 (Note: Because of overlap of collectors into more than one decade, the collectors column totals to more than 144, the actual count of all collectors known for the period.) Lake in 1849-50, and would have known of the important plant collections taken by that individual. It seems probable that the collec- tions were taken earlier than the 1857 date indicated, when Jane would have been 15 or 16 years of age. The exploration of the region later to be circumscribed as Utah was slow. The 1860s was a quiescent period until its final year. In 1869 Sereno Watson arrived on the scene, ac- companied for at least a part of the period by Daniel Cady Eaton (Ewan 1950). The quies- cence of the earlier part of the decade was shattered by the advent of these two, espe- cially by the former. They took some 103 types from the state in that year. Mainly they collected in the Wasatch Mountains of Salt Lake, Summit, and Utah counties. Watson visited the Uinta Mountains and islands of Great Salt Lake in the summer of 1869, and he was in the Raft River Mountains in the northwest during 1868. Eaton evidently ac- companied him into Provo Canyon, where they collected the type of the beautiful scar- let penstemon which was to bear Eaton's name. Watson was associated with the King' survey of the 40th parallel, and he was its principal botanist (Watson 1871). Despite the apparent paucity of their assault on the un- known plant taxa of the state, Watson and Eaton yielded a huge assemblage of mate- rials, and averaged some 51.5 Utah plant taxa per collector. That figure stands as the all- time high in the history of plant collection in Utah. Specimens cited as vouchers for taxa re- ported by Watson and Eaton are numbered in a sequence that corresponds to the species number in the report of the expedition (Wat- son 1871). Because of this practice, the same number will often include specimens from widely disparate localities, i.e., "East Hum- boldt Mts., Nevada and Uinta Mts., Utah." When it has been possible to determine that the Utah portion has been selected as lecto- type, I have cited that as the type, but when the lectotype has been selected from Nevada, I have excluded that name from among our Utah types. Some of the names cited in the checklist might be in error. They are includ- ed here for the sake of completeness. This same problem exists with collections by Edward Palmer and Charles Christopher June 1982 Welsh: Utah Plant Types 135 Parry, whose collections were taken astride the Utah-Arizona border, which was not sur- veyed until later. Collectors in the 1870s include Joseph Ellis Johnson, Edward Palmer, Mrs. Ellen Powell Thompson (sister of John Wesley Powell), Lt. George Montague Wheeler, Captain Francis Marion Bishop, Charles Christopher Parry, Lester Frank Ward, and finally, in 1879, Marcus Eugene Jones, whose appearance on the scene presaged a lifetime as resident bot- anist. Jones was to dominate Utah plant tax- onomy until 1923, a total of 44 years. Palmer and Parry concentrated their efforts in south- western Utah, especially in what is now Washington County. Palmer visited the area twice, in 1870 and 1877, and took a total of 58 specimens later designated as types. He was a collector only, depending on others to name the plants he sampled in Utah. Type collections taken by Parry were mainly from his 1874 trip. Parry was in central Utah in 1875, and again in 1877 (with Palmer) at St. George, but these trips added little to his type list for Utah. The total number of his types is 38, and he was responsible for pub- lishing at least 2 of those. According to Jones (Contr. W. Bot. 17: 3. 1930), Parry was in the St. George vicinity for only about a month. His herbarium is at Iowa State University, and contains nvmierous collections by Palmer along with his own materials (Gabel 1981). Charles Christopher Parry was bom in Ad- mington, Gloucestershire, England, 28 Au- gust 1823, and died at Davenport, Iowa, 20 February 1890. He moved with his family to America in 1832, where they settled in Washington County, New York. He received the degree of Doctor of Medicine from Co- lumbia College in 1846, where he was in- fluenced by Dr. John Torrey, professor at that institution (Preston 1897, Gabel 1981). Parry moved to Davenport, Iowa, in 1846. He practiced medicine for a few months only before turning to his greater interest of bot- any. Following a career exploring the West with various governmental surveys, the pur- suit of natural history led Parry to Utah. He came to Utah on a private trip, not associ- ated with any of the surveys. Contrary to Jones's statement (see above). Parry was in St. George from early April to June of 1874, and then moved north to Cedar City and Beaver. His host in St. George was Joseph Ellis Johnson (Jones 1930, personal correspondence of Parry ISC), pioneer pub- lisher, herbalist, and horticulturalist (Cham- berlain 1950). The following year Parry worked out of Spring Lake (Utah County), where he was the guest of Benjamin Franklin Johnson, brother of Joseph Ellis Johnson. His objective for that year was to obtain a collec- tion of the flora in the vicinity of Mt. Nebo, whose flora he decided was depauperate (Parry 1878, Ewan 1950). Parry was an ac- quaintance of Asa Gray, John Torrey, and George Engelmann. He corresponded with each of them and sent his specimens to them for determination and naming. His contribu- tions to the understanding of Utah plants are significant. Edward Palmer was born at Wilton, Coun- ty Norfolk, England on 12 January 1831 and died on 10 April 1911 in Washington, D.C. (McVaugh 1956). He came to America in 1849 where he settled in Cleveland, Ohio. After attending classes at Homeopathic Col- lege in Cleveland for a few months during the winter of 1856-57, Palmer moved to Highland, Kansas— to practice medicine. He assumed the title of doctor, which he was to carry the remainder of his life. Previously he had been on an expedition to South America, Fig. 1. Marcus Eugene Jones (1897), principal pioneer plant taxonomist in Utah (photo courtesy of the Jones family archivist, Charles Walker). 136 Great Basin Naturalist Vol. 42, No. 2 and was soon involved in other expeditions where he collected plant, animal, and arch- eological materials. In 1870 he arrived by rail at Salt Lake City, where he heard a speech by Brigham Young, was introduced to him, and left for southwestern Utah carrying a let- ter of introduction provided by Brigham Young. He stayed in St. George and vicinity for about 10 days prior to departing with a party delivering a threshing machine to St. Thomas, Nevada. They left St. George on 17 June 1870, and spent at least a day and a half beyond that date still in Utah. He noted the grand sight of the Joshua tree forest on the Beaverdam Slope. His total collections in Utah for that year is thought to be very small, for he took only about 200 plants total for the latter part of 1869 and all of 1870 (McVaugh 1956). Only seven of the 58 Utah types taken by him were collected in 1870. Palmer was an acquaintance of C. C. Par- ry, and Parry helped with distribution of the Palmer material. In 1876 Palmer was em- ployed by the Peabody Museum to do some archeological excavations in southern Utah. He was at St. George from December 1876 to June of 1877. During June he moved his base of operations to Red Creek (Paragonah) where he worked in Bear Valley in early July, then on to Beaver, and finally to Spring Lake. The number of plants taken in Utah in 1877 numbered around 500 (McVaugh 1956), and it was from this series that the bulk of his Utah types were named. Palmer depended on others to name his plants. Mrs. Thompson, Captain Bishop, and Les- ter Ward were associated with the Powell surveys of the early-day geological survey. They worked in southwestern, south central, and central Utah. Mrs. Thompson lived for a while at Kanab in 1872, and her collections are from that period. Bishop's collections are from 1872 and 1873, but with little if any provenience data. Some materials credited to Mrs. Thompson obviously came from some distance away from Kanab and were prob- ably taken by members of the Powell survey, including her husband, Almon Harris Thompson. Bishop was born in New York. He enlisted in the Union army at the age of 17. He was made second lieutenant at the battle of Bull Rim first lieutenant at the battle of Antietam, and he was badly wounded at the battle of Fredericksburg. He was discharged with the rank of captain, a title which he carried for the remainder of his life. After the war he at- tended Illinois Wesleyan University, where one of his instructors was Major John Wesley Powell, whose second expedition he joined as topographer. Following his work with the Powell survey, Bishop joined the faculty of the University of Deseret as professor of sci- ence in 1873, where he served until 1877 (Chamberlain 1950). The delicately beautiful Astragalus episcopus Wats, is named in his honor. Ward evidently centered his efforts in the vicinity of the Aquarius Plateau and Rabbit Valley, working out of the hamlet of Glen- wood, Sevier County. The handsome milk- vetch with diaphanous inflated pods. Astra- galus wardii, was named in his honor by Gray, and the beautiful low twinpod, Les- querella wardii, was named by Watson. His locality information is much better than pro- vided by any of his contemporaries, bu t there has been evident confusion asssociated with his labels (Pennell 1920). He was a man of many interests, with expertise in geology, pa- leobotany, botany, and sociology. Ward pub- lished papers in all of those subjects. For a time he was a clerk and librarian at the U.S. Bureau of Statistics. His collections in Utah are from the vicinity of Glenwood, Twelve Mile Canyon, Rabbit Valley, Dirty Devil River, The Button, and Aquarius Plateau (Ewan 1950). Others who collected type materials in Utah in the 1870s were Lt. George M. Whee- ler, Andrew Lafayette Siler, and Townshend Stith Brandegee. The collections of Wheeler are treated, at least in part, in his publication in the United States Geographical Surveys west of the 100th Meridian (Wheeler 1878). Wheeler had little sense of history, as in- dicated in the total lack of geographical in- formation about plants he collected. Appar- ently he took some plants in Salt Lake County, as indicated in a letter by Marcus E. Jones, dated "8. 1. 1904" (on file at BRY), in which he intimated that Wheeler, too, had collected CoUomia dehilis in Salt Lake Coun- ty, but without recording the locality. Bran- degee worked with one of the government surveys, mainly in present-day San Juan June 1982 Welsh: Utah Plant Types 137 County. Siler, a Mormon rancher, lived at a place called "Ranch" near Kanab and took specimens in Kane and adjacent Mohave (in Arizona) coimties. He was much interested in cacti and sent a collection of a low de- pressed-hemispheric ball-cactus to Engel- mann, which was later named by Britton and Rose as Utahia sileri. Jones (Contr. W. Bot. 16: 46-47. 1930) says of Siler: "He was about seven feet tall; and slim as a rail, and wore about a No. 14 shoe. He was awkward and uncouth, but a real man." Unfortimately, the type of the genus commemorating the state of Utah was collected in the vicinity of Pipe Springs, in Arizona, and that generic name is now synonymized within an expanded Pedio- cactus. Siler's collections are mainly at GH and MO (Ewan 1950). The 1880s were relatively quiescent when compared to the efforts and successes record- ed in the statistics of the 1870s. Together, M. E. Jones, Fred Eugene Leonard, Samuel Mills Tracy (alone and with someone named Evans), and William Trelease collected some 54 types. Jones worked in Washington Coun- ty in 1880, and widely in Utah in later years of the decade. Leonard took specimens in northern Utah, especially in Salt Lake Comi- ty. His collections were sent east, where they were named later by Rydberg. William Tre- lease stopped briefly at Helper in 1889, in transit to St. Louis, Missouri, after spending the summer with the Harriman Alaska Expe- dition. He collected the type of a yucca that he named in honor of Mrs. Harriman. Jones moved to Utah in 1880, to Salt Lake City, where he was to reside imtil 1923. At first he taught in the old Salt Lake Academy, a congregational school. From 1881 to 1885 he conducted a private high school. He was one of the early members of the Utah Natural History Society, of which he was president in 1899, and a fellow of the AAAS. Jones was cm'ator of the museum and librarian at the University of Utah for the year 1889-90. His large herbarium of western plants was stored for a time at the University of Utah (Cham- berlain 1950), and it was from there that the collection was retrieved by Dr. P. A. Munz and taken to Pomona College in the 1920s (Richard Shaw, personal communication 1982). The greatest era for collection of Utah plant types occurred in the 1890s. Marcus E. Jones was the main contributor of type mate- rials. Of the 259 types taken during the dec- ade, Jones was responsible for 193 of them. Jones was employed in 1894 by the U.S. De- partment of Agriculture as "Special Field Agent" (Jones 1895). He was able to spend the entire field season collecting plants. Ac- cording to his own account (see Leafl. W. Bot. 10: 208-217. 1965), Jones outfitted a team and buggy that he sent south in early March with a young man as driver. The driv- er and outfit proceeded to Holden, where they were trapped by a late winter storm. Jones traveled by train to Oasis, where he hired a team and reached Holden in a bliz- zard. Jones's account is as follows: "The next morning we drove out into the snow and toil- ed all day in a cloudless sky with the sun daz- zling in our faces and by night our faces were burnt to a blister and the serum was oozing off our chins. That night we camped on the desert in a foot of spotless snow where there was hardly enough brush to make a fire and no water but snow. A better bed I never slept Fig. 2. Alice Eastwood and John Thomas Howell (1936), near Yakima Park, Mt. Ranier, contributors to understanding of botany in Utah and the West (photo by Edith Hardin English, courtesy of John Thomas Howell). 138 Great Basin Naturalist Vol. 42, No. 2 on. The next morning it was bitterly cold to get out of bed and get a fire started in that snow, but warm tea soon thawed us out. That night we slept in mud near Black Rock and the next day were at Milford. Our faces were now all scabs and sores and we looked as though we were suffering from some loath- some disease." Jones was to be in the field, except for short pauses in Salt Lake City, un- til the 18th of October. During the 1894 field season he collected in Washington, Kane, Garfield, Piute, Sevier, Emery, Carbon (then a part of Emery County), Utah, Sanpete, and Wayne counties. His collections amounted to 4060 numbers, a total of about 35,000 speci- mens. Never again would anyone have such an opportunity. The next most important collector of the decade was Alice Eastwood. Born in Canada in 1859, she moved to Denver, Colorado, in 1880, where she taught high school for sever- al years. She was able to devote her life to tlie study of plants by living initially on the proceeds gained through investment. In 1892 she traveled from Thompson's Springs (pres- ent day Tliompson) in Grand County south to San Juan County. A second trip was under- taken from Mancos, Colorado, west to Wil- low Creek in San Juan County, Utah, in the ^:^> Fig. 3. John Torrey (1868), author with John Charles Fremont and Asa Gray of specimens taken in Utah bv Fremont and by Captain Howard Stansbury, teacher of Charles Christopher Parry, and correspondent of Parry, Palmer, and others (photo courtesy of Richard \V. Pohl). svmimer of 1895. These trips allowed her ac- cess to some of the most remote and rugged country in the southeastern portion of the state. She was able to collect numerous plants on both occasions, among them the types of some of our most unusual plant taxa. Her in- terest in the region was to continue until her death in 1953 a tthe age of 94. Her final trip to the state (with John Thomas Howell) was in 1941. Type collections taken by her in Utah span 50 years, longer than for any other contributor. On at leas t the 1895 expedition she was accompanied by B. Alfred Wetherill, famous for his part in the discovery of the Pueblo dwellings at Mesa Verde, and later pioneer archaeologist in the region. He was to send some few things to Miss Eastwood, and this accounts for his inclusion on the list of Utah plant type collectors. Two other important figures in Utah plant taxomomy appear among the cast of collec- tors in the 1890s. They are Per Axel Rydberg, a young immigrant from Westergoethland, Sweden, and Aven Nelson, natural history professor at the infant University of Wyom- ing. Rydberg had arrived in the United States in 1882 and was about to complete his docto- rate at the University of Nebraska, where he wrote a conservative treatment of the Resales of Nebraska. Later he would contribute nu- merous critical publications of the Rocky Mountain flora, and would have an over- whelming role in the botany of the West. His trip to Utah in 1895 seems not to have been long, and he picked only a few things, mainly in the vicinity of Logan. More will be said of him in later parts of this paper. Nelson had only recently arrived at Laramie (in 1887), where he had accepted a position to teach English, but upon his arrival there found that another English teacher had been hired. He was offered the chair in natural history in- stead, and was to contribute to knowledge of western American botany for more than five decades. Completing the list of collectors of types for the decade were Carl Albert Purpus (a German), James Henry Linford (a Utah na- tive), and A. Isabel Mulford, a student of William Trelease at the Missouri Botanical Garden (Ewan 1950). Purpus collected in the La Sal Mountains and in southwestern Utah. Linford's collections are from the northern June 1982 Welsh: Utah Plant Types 139 Wasatch Mountains, chiefly from Logan Canyon, and the place of deposition is un- known. Mulford traveled through the central and northern regions. The period from 1900 to 1909 was marked by a decline in types collected. The total number taken was 105, a large number by modern standards, but low when compared to the previous decade. Too, the number of contributors increased. Jones continued to be active, but what does one do for an encore following such a performance as his of the 1890s? Jones took only 17 types, and Rydberg (alone or with Carlton) accounted for 24 types. That Jones was active is indicated by his collections, which yielded type materials for the years 1900, 1903, 1906, 1908, and 1909. Rydberg's contributions were taken in 1905 from Salt Lake Coimty and from the Tushar Moimtains, astride the Piute-Beaver coimties boundary. The other most important collectors of this time, as regards historical perspective, include Ivar Tidestrom, who was to publish a flora of Utah and Nevada, and Albert Osbun Garrett, resident Utah botanist and teacher for more than four decades. Su- san Stokes (who later treated Eriogonum), Grove Karl Gilbert (?), Leslie Newton Goodding (student of Aven Nelson), Louis Hermann Pammel and R. E. Blackwood, S. J. (?) Harkness, Frederick Vernon Coville (along with Tidestrom), Carleton Roy Ball (of willow fame), and Charles Piper Smith (pro- ducer of infamous works in Lupinits) each contributed one or more types during the decade. The 60 types taken from 1910 to 1919 rep- resented a further decline, emphasized even more by the efforts of 13 collectors for this period. Rydberg, alone and with Albert Os- bun Garrett, worked in Salt Lake, Grand, and San Juan counties in 1911. Their work in the southeastern portion of Utah was the first serious attempt at imderstanding the flora of that region following the pioneer work of Alice Eastwood. Their collections were rich in type materials. C. P. Smith, A. O. Garrett, Willard Webster Eggleston, E. P. Walker, Thomas Henry Kearney and Homer LeRoy Shantz, Edwin Blake Payson, Francis Whit- tier Pennell, Malmsten, Ricker, and Alice Eastwood provided type material during this time. The 1920s were a time of transition. Utah was involved in the consequences resulting from World War I, and the field botanists of the early part of the century were involved in activities that did not include active field investigations. Jones sold his collection to Po- mona College in 1923 and left Utah for Cali- fornia, where he resided until his death in 1934. Rydberg was involved in completion of parts of the North American Flora and was to pass from the scene upon his death in 1931. The scenario shifted from these actors, who had played such important roles in dis- covery, to others, most of whom did not oc- cupy the stage for any lengthy period. Only 19 types were added by nine collectors. Jones took his last in 1923, in Zion Canyon, return- ing to the region of his early pursuits in his old age. A. O. Garrett continued to add spec- imens during the period. His contributions have never received the meritorious notice that they deserved, although the herbarium at the University of Utah bears his name. Al- though Garrett founded an herbarium at East High School in Salt Lake City in 1903, his private collection resides at the University of Utah, to which it was given under terms of a bequest following his death in 1948. It is the best early collection extant in Utah. Walter Pace Cottam entered the botanical scene in the 1920s. He graduated from Brig- ham Young University with a Master of Sci- ence degree in 1917 (the first awarded by the university). He returned to the university as a professor in the early part of the 1920s and began a lifetime investigating the state of Utah. His primary emphasis did not M'? with plant taxonomy; he is known for his 'vork in plant ecology. Plants were contributed to the herbarium of Brigham Young University as a biproduct of his investigations of the ecology of the region. He inspired others to follow him into the field and, in fact, has served as an inspiration to generations of scholars. In 1931 he joined the University of Utah and served to found that herbarium also. His early collections are at Brigham Young Uni- versity and the later ones are at the Univer- sity of Utah. The herbarium at the University of Utah was founded in 1870 (Chamberlain 1950), prior to Cottam 's arrival there, but most of the collections had been destroyed. 140 Great Basin Naturalist Vol. 42, No. 2 misplaced, or pilfered by that time. He estab- lished the herbarium on a permanent basis in 1933. He had founded the herbarium at Brig- ham Young University in 1923. The Utah State University herbarium was formalized in 1934. Other collectors during the 1920s were E. B. Payson (botanist at the Uinversity of Wyoming), Charles Piper Smith (professor of botany at Utah State during the period 1909- 12, Ewan 1950), Pilsbry, George Everett Os- terhout (amateur botanist and lumberman, whose herbarium is at RM), Nord & Sargent, Fallas, and Mildred Esther Mathias. Payson worked on revisions of genera important in our flora, i.e., Thelypodiian and its relatives and Cryptantha (as Oreocarya). Mildred E. Mathias treated the complex and difficult genera in the Umbelliferae. With Rydberg and Jones both gone from the scene, the 1930s became a time for re- evaluation of western American plant tax- onomy. The divergent views of taxonomy as exemplified by them were put to the test by no less than 30 individuals who worked in Utah in the decade. Some of these, William D. Stanton (a student of Cottam who worked on tlie flora of the Henry Mountains), A. O. Garrett, Bassett Maguire, Edward H. Gra- ham, Bertrand F. Harrison, Sevile Flowers, and Perry Plummer, made concerted efforts at continuing the exploration of the state. Others were mainly involved in summary re- visions and monographs of plant genera whose members included Utah as a portion of their ranges. Within this category were George Goodman, Charles Leo Hitchcock, Mildred Mathias, Louis Otho Williams, Fred- erick Joseph Hermann, Alice Eastwood and John Thomas Howell, Susan Stokes, E. B. Payson, Howard Baker, Reed Clark Rollins, Cedric Lambert Porter (curator of the her- bariumn at RM for many years), and Susan Delano McKelvey. Minor contributions were made by collectors named Larson, Clawson, Carter, F. A. Barkley and M. J. Reed, and Tompkins. There occurred a shift within the state dur- ing the 1930s. Bassett Maguire had arrived as plant taxonomist at Utah State University (then Utah State Agricultural College) in 1929. Maguire was the first professional plant taxonomist to be employed on a continuing basis by any of the major universities, at least as far as vascular plants were concerned. He was an enthusiastic collector and teacher. His collections within Utah and the surrounding states formed the basis of the herbarium at Utah State. That herbarium became the cen- ter of plant taxonomy in Utah in the 1930s under Maguire's direction, and continued to hold that status under his student and succes- sor, Arthur H. Holmgren. The herbarium at Brigham Young Univer- sity came under the direction of Bertrand F. Harrison following Cottam's move to Salt Lake City. Harrison was trained in plant physiology, but continued to build the herba- rium throughout his tenure at Provo. This was accomplished during a time when he served as chairman of the department of bot- any, taught 15-25 hours of classes per week (in all botanical disciplines), and functioned on university committees. His favorite class was plant taxonomy, and he instilled the same kind of enthusiasm in his students that he had experienced under the direction of Cottam, who had taught him taxonomy in the 1920s. Slow was the growth of the collec- tion in this period of economic depression and difficulties of travel in the remote re- gions of Utah. Nevertheless, both Harrison and Cottam were able to contribute sub- stantially to the understanding of the flora and its ecology. The work of Maguire and his associates clearly places Utah State at the center of tax- onomic study in Utah during the 1940s. Ma- guire, Holmgren, and C. L. Hitchcock (with whom Maguire collaborated), formed a nu- cleus about which Utah taxonomy func- tioned. Other workers, all from outside Utah, collected in the state in the decade. Elzada Clover and Lois Jotter traversed Glen Can- yon in 1940. Alice Eastwood (then in her 83rd year) and J. T. Howell were in Utah in 1941, allowing Miss Eastwood a final view of the scenes of her earlier explorations. Dwight Ripley and the amazing Rupert Charles Barneby (already working on in his revision of the monumental genus Astragalus) visited the state first in 1942. Barneby, an English- man, with field experience in Spain, where he first encountered the genus Astragalus, was inspired to come to North America after seeing the name of Astragalus asclepiadoides June 1982 Welsh: Utah Plant Types 141 in a catalogue of North American plants. He then examined the single specimen taken by Jones on deposit at the herbarium at Kew and made the decision that he must see the plant in vivo. Lyman Benson, expert on Ra- nunculus and cacti, was in the state in 1948 and 1949. Frank Gould, specialist in grasses, collected here in 1941. A person named Marsh took materials in both 1945 and 1946. The 1950s mark a period of quiescence in exploration and collection of Utah plant types. Maguire had left Utah State for the New York Botanical Garden, and the post World War II era found tlie professors in- volved with teaching and other pursuits, which did not allow them to allot much time to collecting. Fewer types were taken in this decade than any time since the 1850s. Arthur Hermann Holmgren, B. F. Harrison, C. L. Porter, Rupert C. Bameby, F. J. Hermann, and others contributed only 13 types in the period. The list gives Utah State and Brigham Yomig about equal, if miimpressive, billing. Collectors during the 1960s were affiliated with Utah State (Arthur Holmgren, his son Noel Hermann Holmgren , and Garrett Da- vidse), New York Botanical Garden (Rupert C. Barneby and Johnnnie L. Gentry), and Brigham Yoimg University (Craig Hanson, Stanley Larson Welsh, Glen Moore, James Reveal, Larry Charles Higgins, Stella Leim- omi Tree Welsh, and Nephi Duane Atwood). Ishan A. Al-Shebaz (of Harvard University) was tlie only other collector of Utah plant types in the decade. Personnel from Utah State and New York Botanical Garden were engaged in a collaborative study of the inter- mountain flora preparatory to publication of a manual of that flora. The school that devel- oped at Brigham Young University resulted from the serendipity that followed the ac- ceptance of the position of plant taxonomist (the first for the university) by Welsh in Sep- tember of 1960. Welsh, his wife, and his stu- dents began a concerted effort at exploration of the flora of Utah. The nucleus of the her- barium developed by Cottam and by Harri- son allowed for continued expansion as the flora of the state was investigated. Explor- ation and collection produced some 50 speci- mens that have been named as types at the present. During the 1970s some 47 type specimens were taken. The essential group of collectors remained largely the same as in the previous decade, but there was some shift in in- stitutional affiliation and new names ap- peared. Noel and Patricia Kern Holmgren became affiliated with New York Botanical Garden, and James Reveal was employed by the University of Maryland. Leila Shultz joined Arthur Holmgren as curator of the herbarium at Utah State University late in the decade. The roster of people at Brigham Young Universty who aided in collection of type materials include S. L. Welsh, N. D. At- wood, Joseph Richard Murdock, Frederick J. Peabody, Kaye Hugie Thome, Kathryn N. Taylor, Elizabeth Janet Chase Neese, Susan White, Kent Ostler and David Anderson, Kim Despain, Betty Jean Welsh, and S. L. T. Welsh. Counties of Collection An examination of types taken on a per county basis is instructive as to where em- phasis was given in early botanical excur- sions. Data on types by county are presented in Table 3. Washington County was and is a favorite site for investigations by those tired of the cold and snow of winter, and, because of the warm desert vegetation, which reflects its climate, many people traveled there to collect. Palmer, Parry, and Jones were there Fig. 4. Daniel Cady Eaton (1864), author of the Com- positae in Watson's Botany, who is honored by the name Penstemon eatonii Gray (photo courtesy of Richard W. Pohl). 142 Great Basin Naturalist Vol. 42, No. 2 early, in the 70s and 80s, when many of the taxa were undescribed. Because of their at- tention to that delightful region for early spring and autumn floras, Washington Coun- ty stands in first position with 163 types. Salt Lake County is second with 102, in part be- cause of its unique flora that extends from moderate to very high elevations, and in part because of its position within the state. Early taxonomists visited Salt Lake City, center of commerce and hub of transportation for this vast region. Access to the mountainous areas around the valley of the Great Salt Lake was relatively easy for those operating from this base. Jones's visit to the Wasatch Mountains in 1879 yielded many types. Watson, Jones, Rydberg, Garrett, Stokes, and many others spent time in the canyons and ranges in close proximity to Salt Lake City. The large num- ber of types from San Juan and adjacent Grand counties is attributable in large part to the early work by Eastwood and Rydberg and Garrett. Garfield County owes its large Table 3. Utah types listed by county of collection. County Number Beaver Box Elder Cache Carbon Daggett Davis Duchesne Emery Garfield Grand Iron Juab Kane Millard Morgan Piute Rich Salt Lake San Juan Sanpete Sevier Summit Tooele Uintah Utah Wasatch Washington Wayne Weber Unknown Total 25 7 28 21 3 9 19 45 56 43 20 23 59 19 1 35 2 102 85 22 48 53 36 33 30 9 163 26 4 47 1073 compliment of types to the early work by Rydberg, and to later attempts by more re- cent collectors. Summit County became im- portant in types largely, 1 believe, because of the access to the westernmost portion of the Uinta Mountains, and especially because of the early trip to the area by Watson. Tremendous effort and expense is neces- sary for exploration of the state. Most of the remaining endemics, if past experience is cor- rect, will be found in areas that remain to be explored in remote parts of the state. Each year a few more are discovered, and with each new generation of monographers addi- tional types will be designated from among collections of the previous years. Authors of Taxa Based on Utah Types Those who collect types may or may not be prepared to describe taxa based on them. Indeed, few are those who recognize, imme- diately upon collection, that the material col- lected is undescribed. Often, new taxa are made to fit into concepts of previously de- scribed entities, whether they fit well or not. Each worker is acquainted with only a por- tion of the var ktion of the total flora, and few consider themselves sufficiently prepared to name as new all portions of the variation encountered by them. To some investigators any variation is worthy of a name, regardless of the importance of that characteristic in the total range of variation available. To them the value of a single character is over- riding—the character makes the species, as Linnaeus stated. Because of differences in training, philosophy, and expertise, and others due to change in rmderstanding over more than a century of time, the lasting qual- ity of names recognized as belonging to vi- able entities varies from author to author, and from group to group treated by the same author. There have always been more spe- cialists involved with naming than with col- lecting. Beacuse of this, many authors never saw the taxa that they named in vivo— they were herbarium botanists with experience in dried and pressed plants only. Among the herbarium botanists quality of judgments var- ied as much as between them and their field- experienced counterparts. Gray and Watson represent the two extremes, and both were June 1982 Welsh: Utah Plant Types 143 successful descriptive botanists whose named taxa have stood the test of time. The syn- drome of "name in print at any cost" af- flicted only a few of the major authors (i.e., Edward L. Greene and Charles Piper Smith). The names of authors of Utah types are listed in Table 4. The decade of publication of Utah types, number of authors, and average nmnber of names per author are cited in Table 5. There has always been a disparity among botanists regarding species concept and ap- plication of names. As there has always been a problem of distinguishing between no- menclature and taxonomy, a problem has also existed between botanists in the recognition of species in which infraspecific categories are allowed. Some have chosen to draw spe- cies concepts that are broad and allow for the inclusion of several infraspecific taxa in a hi- erarchial system. Others have felt that such a nested system, in which species, subspecies, varieties, subvarieties, and forms were al- lowed, represented a return to pre-Linnaean phrase names. To them such usage was and is intolerable; the binomial system of nomencla- ture should be adhered to at all costs. When this concept is followed, and when it is coupled with the concept that any variant is worthy of a name, it is possible to multiply names inordinately. Names based on Utah types were supplied by workers from both of the divergent schools, and by others between. Still other workers were not bound by the sense of logic provided by either school; they were self- taught and were trapped in their pre- conceived notions that any variation con- stituted a case for a new species, and that the species were immutable. Because of the dif- ferences in approach, in imderstanding, and in training, our Utah plants were given a va- riety of treatments. The variation among bot- anists today is not as great, but there is still variation. A current move has been to recog- nize only one infraspecific category, the sub- species. If such a move were to succeed, then all variants now recognized at other in- fraspecific levels would be treated as sub- species, and levels of variation would be hid- den by the apparent uniformity of the category. It is true, of course, that taxonomy should not be dependent upon nomenclature. but where the disciplines can supplement each other that course seems best to this worker. Some of the cast of authors is the same as for the list of collectors, but many are differ- ent. Noted is the fact that some of the collec- tors of many types failed to publish new taxa based on their own collections. That task fell to others, either by default or on purpose. One will note that Jones named only 167 taxa but collected some 305. The others were named by some 44 botanists during and sub- sequent to Jones's lifetime, the latest by Re- veal (1977). This was particularly disturbing to Jones, not because others purloined his material, but because he did not feel that most of them were worthy of being named. The 1840s demonstrate only two authors of Utah taxa, i.e., John Torrey and John Charles Fremont, and John Torrey and Asa Gray. Torrey was situated at Columbia College, New York, and Fremont was then engaged in explorations of the West. Later, Fremont would be a candidate for the presidency of the United States (in 1856). Together they named Perchiton occidentale (Torrey & Fre- mont 1845). Torrey and Asa Gray collabo- rated in naming Gilia stenothyrsa and Se- necio multilobatus (Torrey and Gray 1849). All these taxa are based on specimens taken by Fremont. Asa Gray, working out of Fig. 5. Sereno Watson (ca 1870), collector and author of Utah plant types, second in importance only to Mar- cus E. Jones (photo courtesy of Richard W. Pohl). 144 Great Basin Naturalist Vol. 42, No. 2 Harvard University, was the principal 19th- century American plant taxonomist; his con- tributions are legion. Five authors described taxa based on Utah types in the 1850s. These include John Tor- rey, Torrey and Gray, Elias Durand, George Bentham, and Edward Griffin Beckwith. Their published taxa are based on collections by Stansbury, Creuzfeldt, Beckwith, and Car- rington. Seven of the 12 taxa stand at some taxonomic level in contemporary treatments. Perhaps the best known of the taxa described in the decade is Phaca mollissima var. uta- heivsis named by Torrey and now cited as As- tragalus utahensis (Torr.) T. & G. Torrey and Gray were the principal American tax- onomists of the period. John Torrey (1796-1873) was a professor at Columbia College. He became associated with Asa Gray (1810-1888), his student, in writing a "Flora of North America" (1838-1840). Gray Table 4 continued. Author Number of names Years active Table 4. Authors of names of Utah plant types. Author Number of names Years active Al-Shehbaz, I.A. (b.?) Anderson, L.C. (13-1936) Atwood, N.D. (b.l938) Bailev, L.H. (1858-1954) Baker, M.S. (1868-1961) Baker, M.S. (1868-1961) &J.C. Clausen (b. 1891) Ball, C.R. (1873-1958) Barkley, T.M.(b. 19.34) Bameby, B.C. (b.l911) Barneby, B.C. (b. 1911) & N.H. Holmgren (b. 1937) Beal, W.J. (18a3-1924) Beaman, J.H. (b.l929) Benson, L.D. (b.l909) Bentham, G. (1800-84) Bicknell, E.P. (1859-1925) Blake, S.F. (1892-1959) Brand, A. (186.3-19.30) Brandegee, M.K. (1844-1920) Brenckle, J.F. (b.l875) & W.P. Cottam (b.l875) Britton, N.L. (18,59-1934) Britton, N.L. (1859-1934 &J.N. Rose (1862-1928) Buchenau, F.G.P. (1831-1906) Candolle, A.L.P.P. (1806-93) Chaudliri, M.N. (b.l932) Clover, E.U.(b. 1897) & M.L. Jotter (b.l914) Cockerell, T.D.A. (1866-1948) Cottam, W.P. (b.l894) 1 1973 1 1981 4 1972-73 1 1884 2 1938-40 1 1949 1 1921 1 1963 22 1942-66 1 1979 1 1896 2 1957 3 1948-69 1 1856 1 1901 7 1922-37 9 1907-11 2 1900 1 1840 2 1904-08 1 1923 1 1890 1 1864 1 1968 1 1941 2 1904 1 1939 Coulter, J. M. (1851-1928) & J.N. Rose (1862-1928) Coville, F.V. (1867-1937) Cronquist, A.J. (b.l919) Darlington, J. (b.l905) Daston, J.S. (b.?) Dempster, L.T. (b.l905) & F. Ehrendorfer(b.l927) Durand, E.M. (1794-1873) Earle, W.H. (b.?) Eastwood, A. (1859-1953) Eaton, D.C. (18.34-1895) Ehrendorfer, F. (b.l929) Engelmann, G. (1809-84) Engler, H.G.A. (1844-1930) & E. Irmscher (1887-1968) Fedde, F.K.G. (187,3-1942) Flous, M.F. (b?) Flowers, S. (1900-1968) Gandoger, M. (1850-1927) Garrett, A.O. (1870-1948) Gates, R.R. (1882-1962) Gentry, J.L. (b.?) Goodding, L.N. (b.l880) Goodman, G.J. (b.l904) &C.L. Hitchcock (b. 1902) Gould, F.W.(b. 191.3) Gray, A. (1810-1888) Greene, E.L. (1843-1915) Greenman, J.M. (1867-1951) Greenman, J.M. (1867-1951) 6f E.M. Roush (b.l886) Hackel, E. (1850-1926) Hanson, C.A. (b.19,35) Heil, K.D. (b.l941) Heimerl, A. (b.l857) Heiser, C.B. (b.l920) Heller, A.A. (1867-1944) Hermann, F.J. (b.l906) Higgins, L.C. (b.l936) Hitchcock, A.S. (1865-19,35) Hitchcock, C.L.(b. 1902) Hitchcock, C.L. (b.l902) & B. Maguire (b.l904) Holm, T.H. (1854-19,32) Holmgren, A.H. (b.l912), L.M. Schultz(b.l946), & T.K. Lowrey (b.?) Holmgren, N.H. (b. 19,37) Holmgren, N.H. (b.l937) & A.H. Holmgren (b. 1912) Holmgren, N.H. (b. 19,37) & P.K. Holmgren Hopkins, M. (b.l906) House, H.D. (1878-1949) Howell, J.T.(b. 1903) Johnston, I.M. (1898-1960) Jones, M.E. (18.52-1934) 81888-1910 3 1892-93 9 194,3-63 1 1934 3 1946 2 1965 2 1860 1 1980 381893-1942 15 1871 2 1956 13 1871-96 1 1916 2 1909-13 1 1934 1 1949 9 1905-20 1 1917 1 1915 1 1979 5 1904 1 1932 1 1942 65 1852-84 441896-1912 5 1914-17 1 1929 1 1896 3 1962 3 1979 1 1902 1 1961 1 1905 2 1934-37 3 1968 1 1933 6 1941-52 1 1947 1 1891 1 1976 8 1973-79 1974 1 1979 1 1937 1 1906 4 1940-43 6 1923-52 171880-1933 June 1982 Table 4 continued. Welsh: Utah Plant Types Table 4 continued. 145 Number Years Author of names active Klein, W. (b.?) 1 1962 Koehne, B.A.E. (1848-1918) 1 1895 Kuntze, C.E.O. (1843-1907) 5 1885-95 Leveille, A.A.H. (1863-1918) 1 1905 Macbride.J.F. (1892-1976) 3 1916 Macbride,J.F. (1892-1976) & E.B. Payson (1893-1927 1 1917 \4ackenzie, K.K. (1877-1934) 1 1917 Maguire, B. (b.l9()4) 18 1941-51 Maguire,B. (11.1904) & A.J. Cronquist (b.l919) 1 1947 Maguire, B.(b. 1904) & A.H.Holmgren (b. 1912) 1 1946 Maguire, B. (b.r904) & R.E. Woodson (1904-1963) 1 1941 Marshall, W.T. 1 1954 Martin, R.F. (b.l910) 1 1940 Mathias, M.E. (b.l906) 2 19,30-32 McKelvey, S.D. (b.l883) 2 1947 Mez, C.C. (1866-1944) 1 1921 Morton, C.V. (1905-1972) 1 1937 Munz, P.A. (1892-1974) 2 1928-31 Murray, E. 1 1980 Neese,'E. (b.l934) & S.L. Welsh (b.l928) 1 1981 Nelson, A. (1859-1952) .32 1900-45 Nelson, A. (1859-1952) & J.F. Macbride (1892-1976) 1 1916 Nelson, E.E. (1876-1949) 21899-1901 Norton, J.B.S. (1872-1966) 1 1899 Osterhout, G.E. (1858-19.37) 1 1926 Ottley, A.M. (b. 1882) 1 1944 Ownbey, G.B. (b.l916) 1 1958 Parry, C.C. (1823-1890) 3 1874-75 Pavson, E.B. (189.3-1927) 5 191.5-26 Pennell, F.W. (1886-1952) 17 1920-37 Petrak, F. (1886-1973) 3 1917 Pilger, R.K.F. (1876-1953) 1 1922 Piper, C.V. (1867-1926) 1 1899 Porter, C.L.(b. 1905) 3 1952 Purpus, J. A. (1860-1932) 1 1909 Raven, P.H.(b. 19.36) 2 1962-69 Rechinger, F. 1 1936 Reveal, J.L. (b.l941) 24 1966-77 Reveal, J.L.(b. 1941) & J.D. Brotherson (b. 19.38) 3 1966-68 Bobbins, J.W. (1801-1879) 1 1871 Robinson, B.L. (1864-1935) 1 1917 Robinson, B.L. (1864-19.35) & J.M. Greenman (1867-1951) 1 1899 Rollins, R.C. (b.l911) 9 19.37-81 Rosendal, CO. (1875-1956), F.K. Butters (1878-1945), & O. Lakela 1 1936 Rydberg, P.A. (1860-1931) 150 1900-29 Schneider, C.K. (1876-1951) 4 1905 Schulz, O.E. (1874-1936) 2 1924-27 Author Number of names Years active Scribner, F.L. (1851-19.38) Scribner, F.L. (1874-19,38) & T.A. Williams (1865-1900) Sharp, W.M. Sheldon, E.P. (b.l869) Shultz, L. (b.l946) & J. Shultz (b.?) Slosson, M.(b.l873) Small, J.K. (1869-1938) Smiley, F.J. (b. 1880) Smith, C.P. (1877-1955) Smith, J.G. (1866-1925) Standley, P.C. (1884-1963) Stockwell, W.P. (1898-1950) Stokes, S.G. (1868-1954) St. John, H. (b.l892) St.-Yves, A. (1855-1933) Suksdorf, W.N. (18.50-19.32) Swallen, J.R. (b.l903) Thurber, G. (1821-1890) Tidestrom, I. (1864-1957) Torrey, J. (1796-1873) Torrey, J. (1796-1873) & J. C. Fremont (181.3-1890) Torrey, J. (1796-1873) & A. Gray (1810-1888) Trelease, W. (1857-1945) Tuzson, J. (b.l870) Underwood, L.M. (185.3-1907) Vasey, G. (1822-1893) Voss, J.W. (b. 1907) Wagner, W.L. Watson, E.E. (1871-19.36) Watson, S. (1826-1892) Watson, T.J. (b.?) Weber, W.A. (b.l918) Welsh, S.L. (b.l928) Welsh, S.L. (b. 1928) & N.D. Atwood (b. 19.38) Welsh, S.L. (b.l928) & R.C. Barneby (b.l911) Welsh, S.L. (b.l928) & S. Goodrich (b.l943) Welsh, S.L. (b.l928) &G. Moore (b. 1917) Welsh, S.L. (b. 1928) &E.J. Neese(b.l934) Welsh, S.L. (b.l928) & J.L. Reveal (b. 1941) Wherry, E.T. (b. 188,5) White, T.G. (1872-1901) Williams, L.O. (b.l908) Williams, T.A. (1865-1900) Wooton & Standley Yuncker, T.G. (b.l891) TOTAL 1911 1899 1935 3 1894 1980 1914 51898-1905 1916 11 1948-51 1899 1916 1940 14 1903-36 1915 1925 2 1931 1931 1871 8 1910-14 9 1850-71 1845 9 1849-70 31896-1907 1 1921 1 1898 2 1876-93 1 1937 1 1981 1 1929 88 1871-88 1 1977 2 1946 18 1970-80 4 1975-81 1981 8 1980-81 1 1963-68 1 1981 6 1968-77 2 1943-44 1 1894 4 1932-37 1 1899 1 ? 1 1960 1073 146 Great Basin Naturalist Vol. 42, No. 2 was to be associated with Torrey throughout his hfe, and in a biography compiled at Tor- rey's death. Gray (1873) noted that ". . . he went again across the continent to California, and . . . enjoyed the rare pleasure of viewing in their native soil, and plucking with his owTi hands, many a flower which he had him- self named and described from dried speci- mens in the herbarium, and in which he felt a kind of paternal interest." The 1860s did not offer much in the way of opportunity for description of Utah plant taxa. Torrey and Gray described Navarretia setosissima, and de Candolle named Quercus stellata var. utahensis, based on a collection presumeably taken by Beckwith "between Salt Lake City and the Sierra Nevada" in 1854. Sereno Watson and Asa Gray dominated the botanical literature dealing with Utah plant types in the 1870s. They were joined by John Torrey, Daniel Cady Eaton, James Wat- son Bobbins, George Vasey, George Engel- mann, Lester Frank Ward, and Charles Christopher Parry in the publication of some 161 taxa. The most important single work of the decade is the report of the U.S. Geologi- cal Survey of the 40th Parallel, Volume 5, Botany, authored by Watson (1871), but in collaboration with D. C. Eaton (honored in the name Penstemon eotonii Gray), George Engelmann, and J. W. Bobbins. The work in- cluded 51 taxa based on Utah types (39 by Table 5. Decade of publication of Utali types, num- ber of authors, and average number of names per author. Number of Number of Decade names authors N/A 1840-49 2 2 1.0 1850-59 14 5 2.8 1860-69 4 3 1.3 1870-79 166 10 16.6 1880-89 27 6 4.5 1890-99 181 24 7.5 1900-09 180 18 10.0 1910-19 145 25 5.8 1920-29 59 19 3.1 1930-39 61 27 2.3 1940-49 75 28 2.7 1950-59 25 11 2.3 1960-69 44 16 2.8 1970-79 56 14 4.0 1980-81 24 12 2.0 Total 1073 167 6.4 Watson, 10 by Eaton, one by Engelmann, and one by Bobbins), mainly collected by Watson and Eaton in 1869. Watson ranks second only to Jones in number of types col- lected, and is third after Jones and Bydberg in the number of taxa described on the basis of Utah type material. The tremendous effort of the 1870s ap- pears to have almost exhausted the botanical authors. The 1880s were, by contrast, rela- tively quiescent. Only 27 taxa established on Utah plants were named in the period. Gray named 13; Watson only two. The remainder were divided among Vernon Bailey, Otto Kuntze, and John Merle Coulter and Joseph Nelson Bose. Marcus E. Jones sent his early collections taken during this period to Gray and Watson, but when he felt unjustly con- trolled by their editorship of some eastern journals, Jones bided his time until Gray was dead (in 1888), before beginning to publish in earnest in the 1890s. If the 1880s were relatively quiet, they were a calm before the storm. The death of Gray in 1888 and of Watson in 1892 allowed for an increase of activity acro.ss the West. The removal of these two giants of American taxonomy from the scene coincided with the coming of age of Jones in his exploration of Utah. His species concepts had been obtained from his early training, when Gray's Manual of Botany (published in 1867) had influenced him. He had received reinforcement in his concept by early contact with Gray but most especially, from his application of earlier perceived concepts with plants as he found them in the field. Jones, with only a meager contribution in the 1880s, named a staggering 118 taxa from Utah in the 1890s, 75 of them in a single publication (Proc. Calif. Acad. II, Vol. 5. 1895). Jones did not occupy the field alone. Some 24 others authored taxa from among the wealth of undescribed botanical materials. The list of authors includes: Ber- nard Adalbert Emil Koehne, Franz George Phillipp Bucheneau, Carl Ernst Otto Kuntze, Herman Teodor Holm, Fredrick Vernon Coville, A. Eastwood, George Vasey, Ed- mund Parry Sheldon, Theodore Greely White, Edward Lee Greene, William James Beal, John Kunkel Small, Eduard Hackel, William Trelease, George Engelmann, Per June 1982 Welsh: Utah Plant Types 147 Axel Rydberg, Thomas Henry Kearney, Lu- cien Marcus Underwood, Aven Nelson, Thomas Albert Williams, Benjamin Lincoln Robinson, Jesse Moore Greenman, Jared Gage Smith, Charles Vancouver Piper, and Elias Emanuel Nelson. Alice Eastwood and Per Axel Rydberg were both beginning their activities within Utah; they would spend dec- ades involved, in part, with the Utah flora. The decade was the golden age of descriptive botany in Utah, and in deference to Jones's contributions the decade should be known as the "Jonesian Era." The decade from 1900 to 1909 showed only a slight decline in taxa named from the state when compared to the previous decade (180 compared to 181), with the deficit in types collected in that period (105) being made up by names being applied to speci- mens taken previously. Some 18 authors as- saulted tlie collections of the state. Jones did not come close to matching his unparalled feats of the 1890s. He named only 21 taxa in the decade. Rydberg, an enthusiastic and energetic professionally trained botanist, Rished to fill whatever vacuum was left fol- lowing Jones's valliant efforts. Introduced to the region in the mid-90s, Rydberg had named only one taxon from Utah prior to the turn of the century. Now he assumed the leading position as author of 65 Utah types in the decade, in preparation for publication of his flora of the Rocky Mountains (1917). Prin- cipal authors besides Rydberg and Jones were Edward L. Greene (21 names) and Aven Nel- son (20 names). Rydberg had no close competition in the period from 1910 to 1919. He named another 65 taxa based on Utah materials, and Jones named only 14. The 1920s were the lowest for publication of new taxa since the 1880s. Only 59 taxa were named by 19 authors. Rydberg authored only 9, second to F. W. Pennell, then engaged in a revision of the genus Penstemon, who named 14 taxa. Neither Jones nor Rydberg were to live beyond the early 1930s. Jones named a final 4 taxa in that decade. The period fell to Sidney Faye Blake (4), Alice Eastwood (4), Ivan Murray Johnston (2), David Daniels Keck (4), Aven Nelson (2), F. W. Pennell (3), and Susan Gabriella Stokes (12), largely without a lead- er. Leadership in the naming of Utah's plants in the 1940s clearly belongs to Bassett Ma- guire, who accepted a position at Utah State Agricultural College in the 1930s. Others working in the flora of the state in the 1940s, who contributed names of new taxa, include Rupert Charles Barneby, Arthur John Cronquist, Alice Eastwood, C. Leo Hitch- cock, John Thomas Howell, Ivan M. John- ston, and David D. Keck. The 1950s were relatively quiet. This peri- od was marked by the shift from efforts of World War II, and Bassett Maguire had left Utah State. Charles Piper Smith, in his senes- cence, and suffering from a lack of under- standing of what constituted a species within Lupinus, named seven inconsequential lu- pines as species, all but one of them from the head of Salina Canyon, and all of those be- longing to Lupinus sericeus in a broad sense. Barneby named five species during this period. In the decade of the 60s Barneby (with 9 new taxa) was displaced by James Lauritz Reveal (with 15 new taxa) as leader in au- thorship of new names. Reveal had begun his assault on Eriogonum, a genus rich in endem- ic taxa and other narrowly restrcted plants. Craig Hansen, James L. Reveal, and Larry K Fig. 6. Asa Gray (ca 1865), author of numerous Utah types, based on collections corresponded by Watson, Parry, Thompson, Siler, Bishop, Palmer, and Jones, and the most important North American plant taxonomist of the 19th century (photo courtesy of Richard VV. Pohl). 148 Great Basin Naturalist Vol. 42, No. 2 Charles Higgins and Stanley L. Welsh, all as- sociated with Brigham Young University, named a third of all species described during the decade. Botanical exploration continued at an ac- celerated pace in the 1970s. Impetus for pub- lication of the taxa discovered in this period was supplied by preparation of the grand plan for publication of the "Intermountain Flora" by members of New York Botanical Garden and associates. Included in that team are Arthur J. Cronquist, Arthur H. Holmgren, Noel H. Holmgren, Patricia K. Holmgren, and James L. Reveal. Noel Holmgren, alone or in combination with others of the group, published descriptions of 10 new taxa from Utah, and Reveal published 1 1 . Work on exploration of floristic provinces of Utah by botanists from Brigham Young Universtiy was expanded in the 1960s, and was again intensified in the 1970s. The Utah Flora project, initiated in the 1960s, offered stimulus. Further activity was increased by passage of environmental protection laws, and especially by passage of the Endangered Species Act of 1973, as amended in 1978. Welsh, alone or with others, described 29 taxa in the decade. That is the largest number involving any individual since the 1910-1919 period. Atwood named 4, bringing the total for those at Brigham Young University to 33, or 61 percent of the total for the decade. The Decades Beyond Tliat the past is prologue is illustrated by the fact that several new taxa have been de- scribed from Utah since the close of the peri- od covered by this text. Those which have been published are included in the list of type material, but they are not summarized as for previous decades. Too, I recognize that a decade does not begin with the zero year, but ends with it. For the sake of convenience on my part, I have chosen to use it otherwise. Those who follow can find fault, as all who view others' work must, but it will not mate- rially affect the data presented here or my view of the history of plant collection in Utah. Perhaps some will find this summary iLseful, and possibly someone will choose to keep it current for the future. Marcus Eugene Jones— His Contributions AND His Contemporaries In the summer of 1879 (Broaddus 1935) an event occurred which, because of its hap- penstance and because of its forthrightness, would not seem to have any lasting effect. A young botanist, on one of his earliest trips west from the rolling farmlands of Iowa, hap- pened to find a wallet (Munz 1965) that be- longed to a General Wm. J. Palmer. The wal- let contained valuable papers and was returned to its owner. Following this act of an honest man. General Palmer sent the bot- anist, Marcus E. Jones, on repeated trips to explore regions for mining or agricultural po- tential. Jones was enabled to travel widely in the western portions of the continent, all be- cause of serendipity— the finding and return of another's property. Because of this, west- ern American plant taxonomy was altered for all time. He indicated his gratitude to Palmer (Jones 1891) by naming Cleomella palmerana; "Dedicated to General Wm. J. Palmer, than whom there is no one more interested in sci- entific researches in Utah, or who has shown more interest in a more substantial way." Jones was born in Jefferson, Ashtabula County, Ohio, on 25 April 1852 and died at San Bernadino, California, in his 83rd year on 3 June 1934. He was reared and educated near Grinnell, Iowa, having moved west with his family in 1865 ("on the very day that Lin- coln was shot "), graduating from Grinnell (Iowa) College in 1875 (Broaddus 1935). In 1876 and 1877 the youthful botanist collect- ed plants in Iowa. Early in 1878 he left for Colorado, where he collected large numbers of specimens that were sold as "sets." He re- turned to Colorado Springs in 1879 as "pro- fessor of natural science." During July he traveled to Salt Lake City and collected at Alta, City Creek Canyon, Lake Point, and Bingham Canyon. Jones moved from Iowa to establish per- manent residence in Salt Lake City in 1880. Accompanying him was his bride (since Feb- ruary 18), Anna Elizabeth Richardson (1853-1916), who would spend most of the remainder of her life with Jones in Salt Lake City. Instructive is his account (Munz 1965) of the itinerary from Salt Lake City to June 1982 Welsh: Utah Plant Types 149 Washington County and return during the spring of 1880. 'T left Iowa Falls, Iowa, Febmary 18th for Salt Lake City and got there the 24th. March 3rd I hired a team and started to drive to St. George. I reached Sandy. On the 4th I re- turned to Salt Lake and back to Sandy (hav- ing forgotten something). On the 5th reached Pleasant Grove, Spanish Fork on the 6th, Mona on the 8th, and Juab on the 9th, Warm Springs near Gimnison on the 10th, Willow Bend below Richfield on the 11th, Elsinore on the 12th, where I stayed three days and went 14 miles up the Clear Creek Canyon on the 18th. Then went seven miles farther on the 19th, then to Pine Creek fifteen miles above Beaver, Beaver on the 20th, Buckhorn Springs on the 22nd, Summit south of Paro- wan on the 23rd, then to Fort Hamilton on the 24th where I found the first spring flow- ers. I reached Leeds on the 25th and St. George on the 26th. I botanized all around the city on the plains and mesas until the 9th of April when I went over on the Santa Clara. On the 14th I went to Washington five miles east. On the 20th I started for Salt Lake and reached Bellevue. On the 21st got to Ka- rarrah (Kanarrah?). On the 22nd Fort Ham- ilton. On the 23rd Parowan. On the 24th. Beaver. On the 27th Com Creek. On the 28 Fillmore. On the 29th to the ridge north of Scipio. On the 30th I reached Nephi. On May 1st I reached Spanish Fork. On the 3rd, I reached the Point of the Mountain and the 5th Salt Lake City." One cannot find in Jones's rather volumi- nous published writings whether his young and beautiful bride of a few weeks accom- panied him on this long and tiring trip. He makes no official acknowledgment that she did go with him. In an account of Anna Jones's life written by her daughter (Broaddus 1952), however, it is noted that "At first Anna went with him on wagon trips of in- credible difficulty and danger, even as far as St. George in southern Utah, helping collect wild flower specimens, pressing and drying them, assembling sets for shipment to Ejro- pean and eastern herbaria, even mounting many of the most attractive flowers in little booklets of 'Western Flora,' which they sold. " Probably she was on this first trip south; it is inconceivable to think otherwise. His respect for women was less than over- whelming. Marcus (called Mark by his family and close friends) wrote of his great admira- tion and love for his mother (Jones and Broaddus 1952), but he demonstrated little regard for other women. His unpublished dia- ry of 1880 (POM) cites his expenses on his wedding day, and then, almost as an after- thought, notes in the margin of the diary, "today I married Anna Elizabeth Richardson." Anna Jones was a talented and educated woman, having taught school for several years from the age of 16, in Iowa Falls, Iowa, and in the surrounding area. She went to Grinell to complete one year of advanced study in English, but remained to complete the college course. Finally, she accepted a position as "Lady Principal" at Grinell (Iowa) College, a position she resigned in January of 1880 so that she could marry one of her former Latin teachers, Marcus Eugene Jones. It is evident from correspondence and from the published account of her life that Anna Jones supported her husband in his life goal of botany. She did sewing for hire, took in individual boarders at first, and finally (1900-1910) established a boarding house to provide funds for Jones to continue his work. Jones had three main goals; to collect the flora of the West, to publish a treatment of Astragalus, and to write a flora of the Great y^WS fS5j Fig. 7. George Engelmann (ca 1864), specialist in cac- ti, yucca, agave, and conifers, and correspondent of Par- ry, Jones, Siler, and others (photo courtesy of Richard W. Pohl). 150 Great Basin Naturalist Vol. 42, No. 2 Basin. All funds provided by Anna and by himself, from mining, consulting, writing, and other projects, were technically supposed to go toward accomplishing of these goals. But Jones was not good at managing money, ex- pending large quantities of capital in photog- raphy and unwise investments. Anna lived to see practically all of the first goal and much of the second one completed. Jones had es- sentially completed the manuscript of Astra- galus by 1910 (Bameby 1964). Neither Anna nor Marcus would live to see the final goal attained. The Great Basin flora would remain for others. Broaddus (1952) notes of her mother that "The last year of her life whe spent in Clare- mont, California, keeping house for her youngest child, Mildred, to help her finish college, and earning what she could by doing dress making." Mildred was one of three chil- dren; the others were Howard and Mabel (Broaddus 1952). f Fig. 8. Albert Osbun Garrett (ca 1930), resident Utah plant taxonomist for more than five decades, teacher, collector, and author of six editions of the Spring Flora of the Wasatch (photo courtesy of Lois Arnow). It is not surprising that Jones was able to accomplish so much, with a wife so dedicated to his care and to his cause. His trip to St. George required 22 days, but he returned to Salt Lake City in just 16 days. This compares with a drive of about seven hours each way today. The state must have seemed overwhelmingly large to Jones, as it still remains, but his youthful enthusiasm seems to have overshadowed any doubts re- garding size or complexity of the flora. In the ensuing decades he researched a sur- prising number of places, including many of the mining areas such as Frisco, Silver Reef, Detroit, and Eureka. The state, despite its great size, did not occupy all his time. He traveled to California, Nevada, Montana, Arizona, and finally to Mexico, but his pre- occupation with Utah plants is evident from his accomplishments. During his 43 years as resident botanist for Utah this resourceful person built a personal herbarium of about 100,000 unmounted sheets (Munz 1965), among them the best set of Utah plants as- sembled in all history. Included among that herbarium were 305 Utah types, almost three-tenths of all types collected in the state between 1843 and 1981. Additionally, Jones had named some 165 plant taxa based on Utah types, mainly on his own collections. If that was not enough, Jones, after becom- ing disgruntled with other publishers (his life is filled with examples of being disgruntled), began to print his own journal "Contribu- tions to Western Botany," on a hand-oper- ated press, using badly worn type, at his home on the avenues in Salt Lake City. This remarkable pioneer undertook to complete a revision of the genus Astragalus for North America. Difficult was the task, at any time, but Jones also faced the problems of trans- portation, communication, and paucity of li- brary materials; and, though largely depen- dent on his own collections, his meager locality data is encyclopedic when compared to that recorded (or unrecorded) by some of his contemporaries. Despite these hurdles, the revision of As- tragalus was essentially completed by 1910 (Barneby 1964). It was not published until 1923, apparently printed as were his "Contri- butions" on his own press, with worn type. The text was distributed on "Feb. 15, 1923" June 1982 Welsh: Utah Plant Types 151 and tlie plates on "June 20, 1923" (Jones 1923). In that same year, Jones's herbarium was piu-chased by Miss Ellen B. Scripps of La Jol- la, California for Pomona College, Clare- mont, California. That herbarium, the richest of any in Jones's first set of Utah type mate- rial, resides today at Rancho Santa Ana Bot- anical Garden, having been integrated there- in in the 1960s. The species concept of Jones was similar to that of Asa Gray, the leading 19th century botanist with whom Jones corresponded, and to whom he sent early sets of his western col- lections. A tribute to Jones abilities is in- dicated in the fact that more than half of the taxa named by him from Utah are still ac- corded taxonomic standing in contemporary treatments. He came to look upon western American plant taxonomy as his own, and summarily defended his territory against all comers. His ego was monumental, but was occasionally tempored by flashes of knowledge that he might not be omnipotent. The field season of 1894 (Jones 1895) found him in the field, hired by the U.S. Department of Agriculture as "Special Field Agent." It was to be his best year, with "about 50,000 specimens and 1700 sets, there being 1106 species and varie- ties in the sets . . ." Or, (Munz 1965) "The to- tal number collected was 4060, and about 35,000 specimens," depending on which of the two accounts of the season one is to be- Ueve. Jones (1895) opened the introductory remarks of the report of the 1894 collections, which is the most remarkable single pub- lication in Utah's history of botanical explor- ation, with the following sentence: "Having had an opportunity to examine the material in the National Herbarium I have been able to correct many errors, and possibly to make a few more." That publication, Jones's "Con- tributions to Western Botany. No. VII.", re- ported on some 119 new taxa for Utah, inter alia. That record will stand for all time. The report was published in "Proceedings of the California Academy (Second Series, Vol. 5)," and was one of the last of his contributions published elsewhere (Jones 1895). There was no one free from Jones's criti- cism, which resulted from a growing para- noia due in part to his isolation from the centers of botanical investigation in the coun- try, and in part to the lack of recognition for his contributions, which are staggering. Nev- ertheless, his criticisms recounted herein were largely unfounded. His contemporaries were, with some exceptions, honorable men doing work in the best tradition open to them. They were, in their own ways, as iso- lated and as unrecognized as Jones. His pub- lished attacks on them yield information and insight into them and him. They are cited here for that reason. Jones thought of himself as benevolent and charitable. The following examples are presented to give an idea of the time in which Jones lived, and to give a portion of the cast of characters who wandered through Jones's existence as a botanist. In his "Contributions to Western Botany. No. 12" (1908), Jones took N. L. Britton to task for provisions of the American Associ- ation for Advancement of Science attempt at botancial nomenclature. One should know that no standardized code was acceptable to the botanists in the United States at that time, and one would not be forthcoming until publication of the International Code of Bot- anical Nomenclature in 1935, following Jones's death. Jones wrote of the Brittonian attempt: "Its chief virtue was in a rigid ad- herence to priority, " and further "... we find that it isn't priority if it goes back farther than 1753, and it is not priority if it is not in the Latin language." And, still further; "If it is contended that common terms are not Lat- in we can easily follow the lead of Britton and put a Latin tail on them (a la Manihota Britton for Manihot). In that case we would have such very attractive combinations as the following: Sagebrushurn tridentatum, Brit- tonastriim tumbleweedum, Greenella slip- penjehna, RydbergioneUa bitterroota, Covillea creosotebusha, Nelsonella greasewooda, etc."l Jones did not agree with acceptance of 1753 as the starting date for botanical no- menclature, and neither did he believe that synonyms should stand in the way of a sub- sequent use of those same names. Latin as an official language for publication of new spe- cies was unacceptable. Jones (1908) wrote: "The purpose of science is to clarify, ... so that those who follow us can take up work where we left off and ultimately bring out all 152 Great Basin Naturalist Vol. 42, No. 2 the underlying facts and explain causes of natural phenomena." Latin, which was in- cluded in his formal education at Grinnell College, and which he had taught in the late 1870s, would not add to clarification for those who would follow him, he thought. Jones carried his criticism to Dr. B. T. But- ler who published a treatment of the western American birches in the Bulletin of the Tor- rey Botanical Club in August of 1909. After systematically chastising Butler on the merits of his work, which criticisms were merited, if not pleasantly couched, he unfairly attacked the education of Butler and the quality of his teachers (Jones 1910). "The writer [Jones] would be glad to subscribe several cents to- ward a fund for the education of the profes- sors who are responsible for this kind of work so that they might go to some botanical kin- dergarten where they could learn that a plant growing in the shade or on a north slope or in a cold lake will differ from another from the same parent which grows in a warmer and drier situation, and that this condition will be reversed when the offspring of these plants occupy reversed conditions; and where they could learn that a poodle dog running around a house is not necessarily two distinct species because they see his tail only on the south side and later on his head only on the north side." Perhaps, but not likely, Jones would have been less harsh on Dr. Butler, but that stu- dent took his Ph.D. degree at Bronx Park and Columbia, always synonymous to Jones with gross misunderstanding of "true" botany. Jones attacked members of the Brittonian school, especially P. A. Rydberg and those as- sociated with him. An enlightening aside on the Brittonian school is found in Jones's (Jones 1913: 27) comments on Gray's Manual, Seventh Edition, in which he states; "One of the most commendable things in the book is the ignoring of the split genera of Britton, Rydberg, and Small. For some years the Brit- tonians have raised a great hue and cry about the new and original work done by them in splitting up Grayan genera. A recent repeti- tion of this is in the Torrey Bulletin where the new Mantial was reviewed through Bronx Park glasses. Anything published by Harvard is like a red flag before a bull to Bronx Park. Whenever Harvard sneezes Bronx Park has a fit, and it is a standing joke among botanists." Jones (1913) then discusses the work by Ryd- berg in Habenaria; "As though Rydberg had done any original work in the genus." And further, "Rydberg tries to justify this work by saying that some of the changes have already been recognized by Europaean botanists. Now if there is any crazy thing that some Eu- ropean botanist has not done in the near or remote past the writer would like to have it mentioned, and to use that as an excuse for doing some other crazy thing is no argument in its favor." The last sentence is as appli- cable today as it was when Jones wrote it. Some contemporary American botanists look to Europe as the wellspring of all that is im- portant or believable in 20th-century tax- onomy. If the Europeans subscribe to splitt- ing then we should follow. The idea is as absurd today as it was in Jones's time; the Europeans have mainly looked at their flora too long, if we have not looked long enough. Jones was not without humor in his criti- cisms. He (1910: 34-42) wrote a rebuttal to a publication by A. A. Heller dealing with no- menclature. Jones wrote: "The publication of the article on nomenclature in my last Con- tributions seems to have given Mr. Heller (in Muhlenbergia) a bad attack of mental colic, recurrent colic. His capacity to appreciate the motive and scope of my criticisms re- minds one of the Englishman who after trav- eling extensively in this coimtry returned home and said to his admiring friends 'The Americans are a very clever people but they have many uncouth expressions, for example they say 'Where am I at', we would say 'Where his my at.' Jones (1910: 35) continued his argument against the principal of priority begun earlier (Jones 1908). After giving other examples of treating names from languages other than Latin, he notes, "Now that he [Heller] has Hellerized all the English names . . . and got- ten the Hellerian tail properly adjusted to all, he will find that Greek antedates the English, then can get up another set of names from the Sanscrit, then the Chinese, then he might take up the Egyptian and give still another batch of new names. By the time the bot- anical public has begun to reccover from the last of these afflictions it is likely that the phonograph and the graphophone will b*^ '"^ June 1982 Welsh: Utah Plant Types 153 perfected that they can take the ripple marks made by sound waves on prehistoric mud (now turned to stone) produced by the inco- herent babbhngs of some of the simian ances- tors of the Brittonians, and reproduce them so that they will be as lucid as some recent descriptions of plants, and will have far stronger claim to priority of publication than the Brittonian checklist had at the time when it was said to have been published. " Jones was wrong to oppose the principal of priority, and because of his obstinacy, he cre- ated synonyms that were unjustified, i.e., the case of Astragalus angustus (Jones) Jones, wherein he refused to use the legitimate re- placement "ceramicus" proposed by Sheldon because earlier names were already occupied. Priority ultimately provides for a stability in nomenclature; without it chaos could result. Jones should have understood the necessity for a consistent nomenclatiire. In the same article in which Jones (1910: 35) attacked Heller, he took the opportunity for a broadside against P. A. Rydberg and E. L. Greene. "Greene and Rydberg especially have been addicted to bolstering up fictitious species and genera by reasoning from analo- gy instead of known facts. Such arguments al- ways get their users into difficulty. I well re- member when it was a stock argument among geologists to claim that the climate of the Arctic was Tropical in the early Qua- ternary because elephants (mastodons) lived there, animals which live only in Tropical climates. Many years afterwards one of these animals was found in a perfect state of pres- ervation imbedded in ice, and it proved to have very long hair and its stomach was full of Arctic plants which it had eaten, showing that it had its home in a frigid climate where that race of animals flourished, and not Trop- ical as has been dogmatically assumed," Jones wrote. The mastodon case is cited by Creationists in contemporary literature as demonstrating existence of tropical climates in high lati- tudes, but the plants taken from frozen stom- ach contents are said to consist of tropical plants., i.e., buttercups for example. The rattle-headed thinking alluded to by Jones, if not altogether warranted in the men he at- tacked, is not attributable to botanists alone. Bameby (1964) has indicated that Jones was wrong in his malicious attacks, especially of Rydberg. There is logic in Rydberg's ap- proach, even if that approach is subject to in- terpretation. He believed in the binomial sys- tem of nomenclature strictly. He (Rydberg 1929) believed that the variation which he observed, and he was a keen observer, should be described. Such an approach called to at- tention those who were his contemporaries and those who followed him. His work was always limited to those few specimens he had available. That he should see only portions of the entire picture is a fault not his alone; all botanists are troubled with the same problem in varying degree. But, it was on Greene and Rydberg that Jones showered his continuing invective. He survived both of them, and heaped in- dignation on them in their later years, and in Greene's case, even following his death. E. L. Greene died on 10 November 1915. Presumeably Jones wrote his scathing obitu- ary of Greene shortly thereafter, but it was Fig. 9. Anna Elizabeth Richardson Jones (1913), wife of Marcus Eugene Jones, who helped him to collect, and who provided money to support him in his botanical works (photo courtesy of Charles Walker). 154 Great Basin Naturalist Vol. 42, No. 2 not published until a decade and a half later (Jones 1929). The opening paragraph of the paper (Jones 1929: 25) entitled "Greene" is as follows. "There have been several notable deaths in the botanical world since my last contributions. Greene, the pest of systematic botany, has gone and relieved us from his botanical drivel. They say that the good that men do lives after them, but the evil is en- terred with their bones. I suspect that his grave must have been a big one to hold it all." Subsequently, Jones (1929) continues; "Greene was first, last and all the time a bot- anical crook, and an unmitigated liar, when it suited him to try to make a point against someone else." Rydberg survived until July of 1931, and had published his monumental Flora of the Rocky Movmtains first in 1917. The work was reissued in revised form in 1922 (Rydberg 1917, 1922). Not until much later did Jones (1929: 9) take note of Rydberg's flora in print. The opening paragraph of Jones's re- view begins: "We are glad that at last Ryd- berg has put his conception of species and genera in book form. His sporadic pub- lications hitherto have shown botanists that he has no conception of either species or gen- era. In addition he accommodates us by saying that any plant deserving of a name should have a specific name. What he really means is that if any fool botanist anywhere in the world has applied a name to a plant as variety fonn or subspecies that name should be raised at once to specific rank, and which he proceeds to do. At the same time he quibbles over priority by position as though it was a matter of any importance. He 'strains at a gnat and swallows a camel.' " "This book is what I have in the past dub- bed bughole botany, species depending on the number of bugholes in the leaves. Ryd- berg has no conception that there is any such science as ecology, or that environment has any effect on species variation. " Jones (1935, published posthumously) wrote an obituary for Rydberg also, in which he was more a gentleman, but the rancor comes through even then. Jones, it seems, never knew when to let an argument end, even when there was no chance for his oppo- nent to speak. Jones (1935: 141) states: "On July 25, 1931 there occurred the second tragedy in American botany, the death of Rydberg before his time, the first tragedy having been the death of J. N. Rose. Their disappointment at the treatment accorded them by the botanical public was, however, partly their own fault; for no one has a right to subordinate his own judgment to that of another, as I think they did to Britton's, for the sake of financial support." Later, Jones (1935: 142) continued, "Rydberg was inclined in his later years to consider himself per- secuted. I never knew that he had any per- sonal enemies. Nor do I think any one ever got after him for personal reasons. I certainly never did, though once or twice I had some reason to feel that he had not treated me fair- ly in private correspondence." Ivar Tidestrom and Aven Nelson were sub- jects of his invective too, and not without some justification. They were troubled by a lack of material to make definitive taxonomic decisions, and they lacked the experience and understanding that Jones had accumulated in his decades of active field and herbarium studies. Despite the quality of their work, as is apparent in their publications (Tidestrom 1925, Coulter & Nelson 1909), their floras were to stand for decades as the authorities on plant names for much of the Rocky Moun- tain and intermountain regions. The pub- lications were used by generations of botan- ists who were never quite satisfied with keys, descriptions (lacking altogether in the work by Tidestrom), or geographical data. The shame in all of Jones's criticism is that he did not complete a work on the flora of the Basin and Range Province of his own. It, too, would have been flawed, as all works are, but with evidence taken over a long lifetime (more than four decades), it would have served well those later generations on whom was afflicted the work of Tidestrom, and the peripheral work of Coulter & Nelson. Jones's opus magnus laboris, his revision of Astra- galus (1923) is flawed, not by the species con- cept or by the descriptions, but by the rattle- headed polychotomous keys, which are diffi- cult if not impossible to follow, and by his persistence in following his own code of no- menclature. The keys, of which he felt un- justifiably proud (as noted by Bameby 1964), are based on a complex number and letter se- quence, which leaves the user wondering June 1982 Welsh: Utah Plant Types 155 where one goes from any particular place. That his species concept has stood the test of time is indicated again in the work by Barn- eby (1964), who published a revision of the North American species of Astragalus some four decades following the appearance of Jones (1923) work on that incredible genus. Bameby was able to recognize some 60 per- cent of the taxa named in that genus by Jones, but only 38 percent of those named by Rydberg, and most of those at varietal rank. Jones had "seen more of the elephant," as it were, than his contemporaries, and with his abilities of observation and his acute awar- ness of more of it than any other of the peri- od, it is presumed that he would have de- scribed both trvmk and tail properly and have known their approximate locations. His con- temporaries suffered, as do many of our own generation of taxonomists, with a myopia of understanding, and they are caught up in preconceived notions of the importance of plant characteristics. Rydberg let the characteristics of pods in Astragalus, which he likewise monographed (1929), dictate its segregation into more than 30 genera. Carried to its logical conclusion, pod featiu-es would ultimately trap its user into absurdities. The use of pod types led Rydberg to place plants of almost identical vegetative appearance into separate genera (i.e.. Astragalus oophorus Wats. var. caules- cens (Jones) Jones in the genus Phaca and As- tragalus beckwethii T. & G. in the genus Pliacomene). Uniformity in plant taxonomy is a trap laid by taxonomists, because the plants vary in accordance with dictates of genotype as that is influenced by habitat. Too, each taxonomic unit is the result of the genetic pathways that produced it, and each is unique both in pathway and in end result. The system must be made to fit the plants, not the other way aroimd. Still, we allow ourselves to fall into the rut; perhaps we de- mand that we do. In a presentation to the Fourth Inter- national Botanical Congress Rydberg (1929a) outlined his approach to naming of plants. The paper is entitled "Scylla or Charybdis," in recognition of the pitfalls of the descrip- tive taxonomist. Rydberg (1929a: 1539) notes, "I will admit that phytographic work in the last 35 vears has been overdone and I myself can scarcely plead 'not guilty' to such a charge." A portion of the problem of "over description" was recognized as duplication of effort, where one taxonomist did not know that the entity was already described. ". . .the more the taxonomist studies a genus or a fam- ily, the more forms he recognizes, and the better he knows the family, the finer splitter he becomes. Then he either distinguishes more restricted genera, or recognizes in- finitesimal species; or perhaps he recognizes large genera and broad species, but splits these into numerous subordinate categories: subgenera, sections, groups, subspecies, varie- ties, forms, etc., without end." Thus, Rydberg outlined the directions which he recognized at the ends of the spectrum of taxonomic studies. His case was overstated, because not all taxonomists are polarized to either one of the two extremes. Rather, they fall remarkably between, because most taxonomists let real- ity, as they understand it, dictate the course they should follow. It is true that reality is subject to interpretation, and no one can ad- mit to imderstanding all of what is real. But few ignore the information that is available .^v'^t.! Fig. 10. Marcus Eugene Jones (1891i, vmH ! - 'Id equipment and plant press, south of Salt Lake City (pho- to Courtesy of Charles Walker). 156 Great Basin Naturalist Vol. 42, No. 2 to them; their interpretations approach real- ity, more or less. The thesis of Rydberg's speech (1929) re- volves aroimd the publication of the "Phy- logenetic Method in Taxonomy" (Hall and Clements 1923). He (Rydberg 1929: 1540) states, "We have seen the endless and in- finitesimal splitting into categories of the German school and for twenty years have been thoroughly disgusted with their absurd nomenclature of trinomials, quadrinomials, and polynomials. We believed that any form that is worth describing is worth a name and that a binomial is better than any other. But we might have forgotten that there is a Scylla as well as a Charybdis." The defence of Rydberg would have been easier except for his "leaping over the edge" in the direction of recognizing all variants at species level. For the feeling expressed in his paper that he had created the species, and had defended and upheld the creations of others, he states, "For years I have admitted Fig. 11. Charles Christopher Parry (ca 1870), student of John Torrey, correspondent of Asa Gray, Sereno Wat- son, and George Engelmann, and important collector of Utah types in 1874 (photo courtesy of Richard W. Pohl). that I am a splitter, but I have prided myself on being consistent, as, for instance, in splitt- ing some genera so as to make them equiva- lent to other genera in the same family or tribe, but I have just discovered that in order to save already described species of other botanists, even if they have a meager excuse for their existence, I have added some of my own of the same category and made my spe- cies concept very inconsistent." Despite the frailties of his approach, Ryd- berg was a gentleman. He did not attack Jones in the press, as Jones attacked him. Time will serve to judge them both. Edward L. Greene (Ewan 1950) was at- tacked more heartily than any of Jones's other contemporaries, and with more justifi- cation. Of the 44 taxa named by him from within Utah only 3 stand in modern tax- onomic treatments. He did not collect from within the state, and relied on plants taken by others for the basis of his taxonomic deci- sions. Perhaps Jones would have been less harsh in criticism of Greene if he had chosen more substantial variants for naming. But Greene was the product of his background, as was Jones, and that background involved training in the clergy (he was a minister). He seems to have believed in the concept of fix- ity of species, and that any variant was there- fore worthy of a name. He also considered himself to be a linguist and would change names involving compounds of Latin and Greek together, without even knowing about the plants themselves. Time has served to judge him, at least as far as his Utah plant names are concerned, as incompetent. Jones judged him more harshly than that. The career of Marcus Eugene Jones ended suddenly, as noted by his daughter, Mabel Jones Broaddus. In Contributions to Western Botany. No. 18. (1935: 132) she wrote: "At this point on June 3, 1934, my father. Prof. Marcus E. Jones met with a fatal accident. He was returning alone from a day's field trip in the San Bernadino Mts. enjoyed in company with the Samuel B. Parish Bot- anical Society when, at an intersection in San Bernadino, his car was struck in the rear by another car and overturned." Marcus Jones (Jones and Broaddus 1952) was "... a man of unusual ability, strone^ June 1982 Welsh: Utah Plant Types 157 character, and amazing versatility. ... his central interest was always botany." Jones's contributions to the understanding of Utah plant taxonomy are overwhelming. He cannot be equaled or surpassed. Utah Plant Types Abies subalpina Engelm. ex Ward Ainer. Naturalist 10: 555. 1876. Pinaceae. = A. lasiocarpa (Hook.) Nutt. Sanpete (?) Co., Wasatch PL, above Gunnison, Ward sn, 1875 (US!). Abronia argillosa Welsh & Goodrich Great Basin Naturahst 40; 78. 1980. Nyctaginaceae. Grand Co., 6 mi s Cisco, S, E., & M. Welsh 16689, 1978 (BRY!;NY!; US!;UT!). Abronia fallax Heimerl ex Rydb. Bull. Torrey Bot. Club 29: 684. 1902. Nyctaginaceae. = A. elliptica A. Nels. Salt Lake Co., Salt Lake City, Jones 1337, 1879 (US!;NY!;UTC!). Abronia fragrans Nutt. ex Hook. var. pterocarpa Jones Contr. W. Bot. 11: 3. 1903. Nyctaginaceae. = A. elliptica A. Nels. Tooele Co., Cottonwood, near Johnson Pass, Jones sn, 1900 (?). Abronia micrantha Torr. var. pedunculata Jones. Proc. Calif. .\cad. II. 5: 716. 1895. Nyctaginaceae. = Tripterocalyx micrantha (Torr.) Hook. Washington Co., St. George, Jones 5101, 1894 (POM!;US!;BRY!;RM!). Abronia nana Wats. Proc. Amer. Acad. 14: 294. 1879. Nyctaginaceae. Beaver Co., near Beaver City, Palmer 404 12, 1877 (ISC!). Abronia pumila Rydb. Bull. Torrey Bot. Club 29: 683. 1902. Nyctaginaceae. = A. elliptica A. Nels. Emery Co., Emery, Jones 554.5q, 1894 (US!;POM!;NY!). Abronia salsa Rydb. Bull. Torrey Bot. Club 29: 684. 1902. Nyctaginaceae. = A. elliptica A. Nels. Salt Lake Co., Salt Lake City, Watson 965, 1869 (NY!;US!). Abronia turbinata var. marginata Eastw. Proc. Calif. Acad. II, 6: 313. 1896. Nyctaginaceae. = A. fragrans Nutt. San Juan Co., Bartons range, Eastwood sn, 1895 (CAS!). Acerates decumbens var. erecta Durand Trans. Amer. Phil. Soc. 11: 174. 1860. .\sclepiadaceae. = Asclepias as- perula (Decne.) Woodson Salt Lake Co., Salt Lake City, Carrington sn, 1857 (P). Acer kingii Britt. in Britt. & Shafer N. Amer. Trees 656. 1908. .\ceraceae. = A. negundo var. interius (Britt.) Sarg. Wasatch Mts., Watson 216, 1869 (US!). Acer sacharum Marsh, var. trilobum E. Murray Kal- mia 10: 2. 1980. Aceraceae. = A. grandidentatum Nutt. Sanpete Co., Ephraim Canyon, Tidestrom 1003, 1908 (US!). Aconitum divaricatum Rydb. Fl. Rocky Mts. 314, 1062. 1917. Ranunculaceae. = A. columbianum Nutt. in T. & G. Salt Lake Co., City Creek Canyon, Leonard 204182, 1884 (NY!). Aconitum glaberrimum Rydb. Bull. Torrey Bot. Club 29: 151. 1902. Ranunculaceae. = A. columbianum Nutt. in T. & G. Washington (?) Co., (S. Utah) Palmer 11, 1877 (NY!). Actinella biennis Gray Proc. Amer. Acad. 13: 373. 1878. Asteraceae. = Hymenoxys cooperi (Gray) Cock- erell Washington (?) Co., Arizona and Utah, Palmer 260, 1877(US!;ISC!;BRY!). Adiantum rimicola Slosson Bull. Torrey Bot. Club 41: .308. 1914. Polypodiaceae. = A. capillus-veneris var. modestum f. rimicola (Slosson) Fern. San Juan Co, Arm- strong Canyon, Rydberg & Garrett 9423, 1913 (NY!). Agave scaphoidea Greenm. & Roush Ann. Missouri Bot. Card. 16: .391. 1929. Agavaceae. = A. utahensis Engelm. var. utahensis Washington Co., St. George, Pal- mer sn, 1877 (MO?). Agave utahensis Engelm. in Wats. var. utahensis Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 497. 1871. Agavaceae. Washington Co., St. George, Palmer s.n. 1870 (US!). Agoseris caudata Greene Leafl. Bot. Obs. & Grit. 2: 124. 1911. Asteraceae. = A. glauca (Pursh) Raf. var. laciniata (D.C. Eaton) Smiley Sevier Co., Salina Canyon, Jones 5438, 1894 (US!;NY!). Agoseris confinis Greene Leafl. Bot. Obs. & Crit. 2: 124. 1911. .\steraceae. = A. aurantiaca (Hook.) Greene Piute Co., near Marysvale, Jones 5893z, 1894 (US!). Agoseris isomeris Greene Leafl. Bot. Obs. & Crit. 2: 123. 1911. Asteraceae. = A. glauca (Pursh) Raf. Summit Co., Uinta Mts., Goodding 1397, 1902 (US!;RM!;ISC!). Agoseris longirostris Greene Leafl. Bot. Obs. & Crit. 2: 125. 191 1. Asteraceae. = A. aurantiaca (Hook.) Greene Sevier Co., Fish Lake, Jones 5743N, 1894 (US!;POM!). Agoseris taraxacoides Greene Leafl. Bot. Obs. & Crit. 2: 123. 1911. .\steraceae. = A. glauca (Pursh) Raf. Piute Co., near Marysvale, Jones 5372, 1894 (US!). Aletes tenuifolia Coult. & Rose Contr. U.S. Natl. Herb. 7: 108. 1900. Apiaceae. = Musineon lineare (Rydb.) Mathias Cache Co., Rydberg sn, 1895 (US!;RM!;NY!). Allium biceptrum var. utahense Jones Contr. W. Bot. 10: 33. 1902. Liliaceae. = A. biceptrum var. biceptrum Salt Lake Co., City Creek Canyon, Jones 6647, 1898 (POM!). Allium brevistylum Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: ,350. 1871. Liliaceae. Summit Co., Uinta Mts., Watson 1180, 1869 (US!;NY!). Allium cristatum Wats. Proc. Amer. Acad. 14: 232. 1879. Liliaceae. = A. nevadense Wats. Washington Co., St. George, Palmer 454, 1877 (US!;NY!;ISC!). Allium pasgeyi A. & N. Holmgren Brittonia 26: 309. 1974. Liliaceae. Box Elder Co., Howell Valley, Hol- mgren et al. 13125, 1960 (NY!;BRY!;US!;UT!;UTC!). Allium reticulatum var. deserticola Jones Contr. W. Bot. 10: .30. 1902. Liliaceae. = A. macropetalum Rydb. Grand Co., Cisco, Jones sn, 1890 (POM!;US!). Allium tribracteatum Torr. var. diehlii Jones Contr. W. Bot. 10: 18. 1902. Liliaceae. = A. brandegei Wats. Summit Co., Parleys Park, Jones 6590, 1901 (US!;CAS!POM!). Allocarya cognata Greene Pittonia 4: 235. 1901. Bo- raginaceae. = Plagiobothrys scouleri (H. & A.) Johnst. Cache Co., Cache Valley, Mulford sn, 1888 (US!;NDG!). Allocarya orthocarpa Greene Pittonia 3: 109. 1896. Boraginaceae. = Plagiobothrys leptocladus (Greene) Johnst. Cache Co., Mulford sn, 1898 (US!;NDG!). 158 Great Basin Naturalist Vol. 42, No. 2 Ahine pabneri Rydb. Bull. Torrey Bot. Club 39: 315. 1912. Caryophyllaceae. = Stelkiria obtiisa Engelni. Bea- ver Co., Beaver Valley, Palmer 54, 1877 (NY!). Amelanchier ittahensis Koehne Wiss. Bei. Progr. Falk- Realgyni. Berl. 95: 25. 1890. Rosaceae. Washing- ton Co., Bellevlew, Jones 1716, 1880 (US!;POM!;UTC!). Amelanchier titahensis Koehne var. cinerea Goodd- ing Bot. Gaz. 37: 55. 1904. Rosaceae. Washington Co., St. George, Goodding 780, 1902 (RM!). Amsinckia eatotiii Suksd. Werdenda 1: 64. 1931. Bo- raginaceae. = A. retrorsa Suksd. Utali, Eaton 251, 1869 (CAS!). Amsinckia utahensis Suksd. Werdenda 1: 106. 1931. Boraginaceae. = A. tesseUcita Gray Salt Lake Co., Salt Lake City, Vicker sn, 1901 (UC!). Amsonia brevifolia Gray Proc. Amer. Acad. 12: 64. 1876. Apocynaceae. = A. jonesii Woodson Washington (?) Co., Palmer 302, 1877 (NY!). Amsonia eastwoodiana Rydb. Bull. Torrey Bot. Club 40: 465. 1913. Apocynaceae. Grand Co., Moab vicinity, Rydberg & Garrett 8468, 1911 (NY!;RM!;US!). Amsonia latifolia Jones Contr. W. Bot. 12: 50. 1908. Apocynaceae. = A. jonesii Woodson Sevier Co., Monroe Jones 6446, 1899 (CAS!;US!;RM!;POM!;BRY!;NY!). Androstephium breviflorum Wats. Amer. Naturalist 7: 303. 1873. Liliaceae. Syn: Brodiaea paysonii A. Nels. Kane Co., near Kanab, Thompson 59, 1872 (US!). Anemone styhsa A. Nels. Bot. Gaz. 42: 52. 1906. Ra- nunculaceae. = A. multifida Poir. Sevier Co., Fish Lake, Jones 5763, 1894 (US!";BRY!;RM!;POM!;NY!). Angelica dilatata A. Nels. in Coult. & Rose Contr. U.S. Natl. Herb. 12: 446. 1909. Apiaceae. = A. wheeleri Wats. Salt Lake Co., City Creek Canyon, Garrett 2127, 1907 (US!;RM!). Angelica leporina Wats. Proc. Amer. Acad. 12: 252. 1871. Apiaceae. = A. pinnata Wats. Wayne Co., Rabbit Valley, Ward 612, 1875 (US!;NY!). Angelica pinnata Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 126. 1871. Apiaceae. Syn: A. leporina Wats. Summit Co., Uinta Mts. Watson 458, 1869 (US!;NY!). Angelica wheeleri Wats. Amer. Naturalist 7: 301. 1873. Apiaceae. Svn: A. dilatata A. Nels. Northern and Central Utah, Wheeler 1872 (US!). Anotites jonesii Greene Leafl. Bot. Obs. & Grit. 1: 102. 1905. Caryophyllaceae. = Silene menziesii Hook. Utah Co., American Fork Canyon, Jones 1372, 1880 (US!;RM!;POM!;UTC!). Antennaria austromontana E. Nels. Proc. U.S. Natl. Mus. 23: 703. 1901. Asteraceae. = A. alpina var. media (Greene) Jeps. Piute Co., Marysvale, Jones 5522, 1894 (US!). Antennaria dimorpha (Nutt.) T. & G. var. macroce- phala D.C. Eaton Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 186. 1871. Asteraceae. = A. dimorpha (Nutt.) T. & G. Salt Lake Co., Salt Lake City, Watson 654, 1869 (US!). Antennaria obtusata Greene Feddes Repert. 5: 241. 1908. Asteraceae. = A. parvifolia Nutt. Uintah Co., Uinta Mts., Goodding 1209, 1902 (US!). Anticlea vaginata Rydb. Bull. Torrey Bot. Club 39: 108. 1912. Liliaceae. = Zigadenus vaginatus (Rydb.) Macbr. San Juan Co., Natural Bridges, Rydberg & Gar- rett 9407, 1911 (NY!;RM!;US!;BRY!;UT!). Antirrhinum kingii Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 215. 1871. Scrophulariaceae. Tooele Co., Stansbury Island, Watson 767, 1869 (US!). Aquilegia caerulea James var. calcarea Jones Proc. Calif. Acad. II 5: 619. 1895. Ranunculaceae. = A. sco- pulonim var. calcarea (Jones) Munz Garfield Co., above Cannonville, Jones 5312a, 1894 (POM!;US!). Aquilegia depauperata Jones Contr. W. Bot. 8: 1. 1898. Ranunculaceae. = A. flavescens Wats. Utah Co., Provo Canyon, Jones sn, 1896 (POM!;US!). Aquilegia flavescens Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 10. 1871. Ranunculaceae. Syn: A. depauperata Jones Salt Lake (?) Co., Wasatch Mts., Watson 35, 1869 (US!;NY!). Aquilegia flavescens Wats. f. minor Tidestr. Amer. Midi. Naturalist 1: 171. 1910. Ranunculaceae. Sanpete Co., Wasatch Plateau, Tidestrom sn (?). Aquilegia micrantha Eastw. Proc. Calif. Acad. II, 4: 559. 1895. Ranunculaceae. Syn: A. pallens Payson; A. Uthophila Payson; A. rubicunda Tidestr. San Juan Co., near Bluff City, Wetherill sn, 1894 (CAS!;POM!). Aquilegia pallens Payson Bot. Gaz. 60: 375. 1915. Ra- nunculaceae. = A. micrantha Eastw. San Juan Co., LaS- al Creek, Payson 443, 1914 (?). Aquilegia rubicunda Tidestr. Amer. Midi. Naturalist 1: 168. 1910. Ranunculaceae. = A. micrantha Eastw. Emery Co., near Emery, Tidestrom 1418, 1908 (US!). Aquilegia scopulorum Tidestr. Amer. Midi. Naturalist 1: 167. 1910. Ranunculaceae. Sanpete Co., Wasatch Peak, Tidestrom 1788, 1908 (US!). Arabis demissa Greene var. russeola Rollins Rhodora 43: 387. 1941. Brassicaceae. Uintah Co., 18 mi N Vernal, Rollins 1757, 1937 (NY!;US!;CAS!;RM!;BRY!;UTC!). Arabis diehlii Jones Contr. W. Bot. 14: 38. 1912. Brassicaceae. = A. pendulina Greene Beaver Co., Mt. Belknap, Jones sn, 1899 (POM!). Arabis glabra (L.) Bernh. var. furcatipilis Hopkins Rhodora 39: 109. 1937. Brassicaceae. Cache Co., Logan Canyon, Maguire .3437, 1932 (RM!;UTC!). Arabis hirsuta (L.) Scop. var. laevis Tuzon Bericht Frein Verein Pflanzengeog. Syst. Bot. 1919: 43. 1921. Brassicaceae. = A. hirsuta (L.) Scop. Utah Co., Ameri- can Fork Canyon, Jones sn, 1880 (UTC!). Arabis lasiocarpa Rollins Systematic Botany 6: 58. 1981. Brassicaceae. Rich Co., 6 mi w Garden City, R.C. & K.W. Rollins 79307, 1979. (GH). Arabis longirostris Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 73. 1871. Brassicaceae. = Streptanthella longirostris (Wats.) Rydb. Tooele Co., Stansbury Island, Watson 72, 1869 (NY!). Arabis oreophila Rydb. Bull. Torrey Bot. Club 34: 437. 1907. Brassicaceae. = A. hjaUii Wats. Salt Lake Co., Rydberg & Carlton 6678, 1905 (NY!). Arabis pulchra Jones var. duchesnensis Rollins Sys- tematic Botany 6: 59. 1981. Brassicaceae. Duchesne Co., 3.8 mie Duchesne, R.C & K.W. Rollins 79113, 1979 (GH). Arabis pulchra Jones var. pallens Jones Contr. W. Bot. 14: 42. 1912. Brassicaceae. Grand Co., Westwater, Jones sn, 1891 (US!;NY!). Arabis setulosa Greene Leafl. Bot. Obs. & Grit. 2: 81. 1910. Brassicaceae. = A. pendulina Greene Piute Co., Marysvale, Jones 5.330, 1894 (US!;CAS!;POM!;NY!). June 1982 Welsh: Utah Plant Types 159 Arabis stokesiae Rydb. Fl. Rocky Mts. 361, 1062. 1917. Brassicaceae. Syn: A. divaricarpa A. Nels.; = A. confinis Wats. Salt Lake Co., Parleys Canvon, Stokes sn, 1901 (US!;NY!). Aragallus majusculus Greene Proc. Biol. Soc. Wash. 18: 12. 1905. Fabaceae. = Oxtitwpis sericea Nutt. Gar- field Co., Henry Mts., Jones 5674, 1894 (US!;PO.\l!;NY!). Arctoniecon humilis Gov. Proc. Biol. Soc. Washington 7: 67. 1892. Papaveraceae. Washington Co., St. George, Parry sn, 1874 iUS!:NY:;ISG!;NDG!). Arenaria fendJeri (Rydb.) Fern. ssp. brevifolia Ma- guire Madrono 6; 2.3. 1941. Carvophvllaceae. = A. fend- leri var. brevifolia (Magiiire) Magiiire Grand Co., LaSal Mts., Magiiire 17972, 1933 (UTC!). Arermria filiortim Maguire Bull. Torrey Bot. Club 73: 326. 1946. Carvophvllaceae. Iron Co., beach, Navajo Lake, Magiiire 19472, 1940 (\Y!;BRY!). Arenaria hookeri Nutt. in T. & G. var. desertorum Maguire Amer. Midi. Naturalist 46: 506. 1951. Carvo- phyllaceae. Duchesne Co, near Duchesne, Riplev & Barneby8701, 1947 (US!;NY!). Arenaria kingii (Wats.) Jones ssp. plateauensis Ma- guire Bull. Torrey Bot. Club 74: 54. 1947. Carvophvl- laceae. = A. kingii var. plateauensis (Maguire) Reveal Iron Co., Cedar Breaks rim, Maguire 19024, 1940 (US!;NY!;UTC!). Arenaria niittallii Pa.\ var. gracilipes Jones Proc. Ca- hf. Acad. II, 5: 626. 1895. Caryophyllaceae. = A. niit- tallii Pax Piute Co., Brigham ' Peak, Jones .5951, 1894 (US!). Argemone corymbosa Greene ssp. arenicola G. B. Ownbey Mem. Torrey Bot. Club 21(1): 118. 1958. Pa- paveraceae. Emerv Co., 37.9 mi s\v Green River, Own- bey 2146, 1954 (US!:CAS!;RM!:BRY!). Arnica arachtwidea Rydb. N. Amer. Fl. 34: 353. 1927. .\steraceae. = A. mollis Hook. Salt Lake Co., Big Cottonwood Canvon, Rvdberg & Carlton 6609, 1905 (NY!). Arnica catidata Rydb. Bull. Torrev Bot. Club 37: 463. 1910. .\steraceae. = A. mollis Hook. Salt Lake Co., Big Cottonwood Canyon. Garrett 1547, 1894 (RM!;NY!). Arnica chamissonis Less. var. longinodosa A. Nels. Bot. Gaz. 30: 199. 1900. Asteraceae. = A. mollis Hook. Piute Co., Marvsvale, Jones 5883, 1894 (RM!;NY!l Arnica jonesii Rydb. Fl. Rockv Mts. 979. 1917. .\ste- raceae. = A. latifolia Bong. Salt Lake Co., Alta, Jones 119, 1879 (NYI;POM!;UTCl;NDG!). Arnica longifolia D.G. Eaton in Wats. Rep. U.S. Geol. E.xplor. 40th Parallel, Bot. 5: 186. 1871. .Aste- raceae. Summit Co., Uinta Mts., Watson 655, 1869 (NY!). Arnica ovata Greene Pittonia 4: 161. 1900. Aste- raceae. = A. mollis Hook. Salt Lake Co., Alta, Jones 1128, 1879 (POM!:NY!;BRY!;UTC!;NDG!). Artemisia norvegica Fries var. piceetorum Welsh & Goodrich in Welsh Brittonia .33: 295. 1981. Asteraceae. Duchesne Co., Garfield Basin, Welsh et al 18970, 1979 (BRY!:NY!). Asclepias itivohicrata Engelm. var. tomentosa Eastw. Zoe 4: 120. 1893. Asclepiadaceae. = A. macrospenua Eastw. Grand Co., Courthouse Wash, Eastwood sn, 1892 (CAS!;NY!). Asclepias labriformis Jones Proc. Calif. Acad. II, 5: 708. 1895. Asclepiadaceae. Wavne Co., Capitol Wash, Jones .5650, 1894 (P0M!;US!;RM'!;BRY!;NY!). Asclepias Jeucophylla Engelm. in Parry Amer. Natu- ralist 9: .348. 1875. Asclepiadaceae. = A. erosa Torr. Washington Co., near St. George, Parry 207, 1874 (MO). Asclepias ruthiae Maguire & Woodson Ann. Missouri Bot. Card. 28: 245. 1941. Asclepiadaceae. Emery Co., Calf Spring Canvon, Maguire 18310, 1940 (US!;Ufc!). Asclepias welshii N. & P. Holmgren Brittonia 31: 110. 1979. .\sclepiadaceae. Kane Co., Coral Pink Sand Dunes, N. & P. Holmgren 9009. 1978 (NY!;BRY!;UTC!). Astephanus utahensis Ejiglem. in Parry Amer. Natu- ralist 9: 349. 1875. Asclepiadaceae. = Cynanchum uta- hense (Engelm.) Woodson Washington Co., near St. George, Parry 209, 1874 (ISC!). Aster canescens Pursh var. aristatus Eastw. Proc. Ca- lif. Acad. II. 6: 296. 1896. Asteraceae. = Machaeran- thera canescens (Pursh) Grav San Juan Co., Willow Creek, Eastwood 45, 1895 (CAS!). Aster glancodes Blake ssp. pulcher Blake Proc. Biol. Soc. W'ashington 35: 174. 1922. Asteraceae. = A. glanc- odes Blake var. pulcher (Blake) Kearnev & Peebles Kane Co., Elk Ranch, Jones 6037, 1894 (US!;RM!;POM!). Aster glaucus var. wasatchensis Jones Proc. Calif. .\cad. II. 5: 694. 1895. Asteraceae. = A. wasatchensis Jones Blake Piute Co., near Marvsvale, Jones 5861, 1894 (US!;RM!:BRY!;POM!;NY!). Aster halophilus Greene Leafl. Bot. Obs. & Grit. 2: 8. 1909. Asteraceae. = A. chiletisis ssp. adscendens (Lindl.) Cronq. Salt Lake Co., Beck's Hot Springs, Garrett 2057, 1906 (UT!;NdDG!). Aster kingii D.C. Eaton in Wats. Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot. 5: 141. 1871. Asteraceae. = Machaeranthera kingii (D.C. Eaton) Cronq. & Keck. Salt Lake Co., Cottonwood Canvon, Watson 507, 1869 (US!;NY!). Aster leucopsis Greene Leafl. Bot. Obs. & Crit. 2: 8. 1909. Asteraceae. = A. chilensis ssp. adscendens (Lindl.) Cronq. Salt Lake Co., Salt Lake Citv, Garrett 1694, 1905 (UT!;NDG!). Aster thermalis Jones Proc. Calif. Acad. II, 5: 694. 1895. Asteraceae. = A. pauciflorus Nutt. Sevier Co., Monroe, Jones 5410, 1894 (POM!;US!). Aster venustus Jones Zoe 2: 247. 1891. .\steraceae. = Machaeranthera venusta (Jones) Cronq. & Keck. Grand Co., Cisco, Jones sn, 1890 (POM!:US!;C.\S!; RM!;BRY!;NY!). Astragalus amphioxys Gray var. cymbellus Jones Rev. Astragalus 215. 1923. Fabaceae. = A. cymboides Jones Emery Co., San Rafael Swell, Jones sn, 1914 (POM!). Astragalus ampullarius Wats. Amer. Naturalist 7: 300. 1873. Fabaceae. Kane Co., Kanab, Thompson sn, 1872 (GH;US!). Astragalus araneosus Sheld. Bull. Geol. & Nat. Hist. Surv. Minnesota 9: 170. 1894. Fabaceae. = A. lentigi- nosus Dougl. var. araneosus (Sheld.) Barneby Beaver Co., Frisco, Jones 1807, 1880 (US!;RM!;BRY!; POM!:NY!;UT!;UTC!). Astragalus argillosus Jones Zoe 2: 241. 1891. Fa- baceae. = A. flavus Nutt. var. argillosus (Jones) Barneby Emerv Co., Greene River, Jones sn, 1890 (POM!; US!;B'rY!). Astragalus argophyllus Nutt. var. cnicensis Jones Rev. Astragalus 207. 1923. Fabaceae. = A. argophyllus var. martinii Jones Utah Co., Thistle, Jones sn, 1898 (POM!). 160 Great Basin Naturalist Vol. 42, No. 2 Astragalus arietinus Jones Proc. Calif. Acad. II. 5: 653. 1895. Fabaceae. = A. cibarius Sheld. Sanpete Co., Fairview, Jones 5544o, 1894 (POM!). Astragalus asclepiadoides Jones Zoe 2: 238. 1891. Fa- baceae. Grand Co., Cisco, Jones sn, 1889 (POM!). Astragalus beckwithii T. & G. var. beckwithii Rep. U.S. Explor. & Surv. R.R. Pacific 2: 120. 1855. Fabaceae. Tooele Co.?, west of Lone Rock, Beckwith, 1854 (GH). Astragalus beckwithii T. & G. var. purpureus Jones Zoe 3: 288. 1893. Fabaceae. Juab Co., Deep Creek Mts., Jones sn, 1891 (US!;POM!;BRY!;NY!). Astragalus castaneiformis var. consobrinus Barneby Amer. Midi. Naturalist 41: 496. 1949. Fabaceae. = A. consobrinus (Barneby) Welsh Wayne Co., Bicknell, Rip- ley & Barneby 8605, 1947 (CAS!;RM!;NY!;UTC!). Astragalus chamaeleuce Gray in Ives var. pan- guicensis Jones Proc. Calif. Acad. II. 5: 671. 1895. Fa- baceae. = A. argophyllus var. panguicensis (Jones) Jones Garfield Co., Pangiiitch Lake, Jones 6023f, 1894 (POM!). Astragalus chloodes Barneby Leafl. W. Bot. 5: 6. 1947. Fabaceae. Uintah Co., 6 mi se Jensen, Ripley & Barneby 7797, 1946 (CAS!;RM!;NY!;UTC!). Astragalus cibarius Sheld. Bull. Geol. & Nat. Hist. Surv. Minnesota 9: 149. 1894. Fabaceae. Syn: A. ariet- inus Jones Utah Co., Utah Valley, Jones 1679, 1880 (MINN;POM!;US!;NY!). Astragalus cicadae var. laccoliticus Jones Proc. Calif. Acad. II. 5; 672. 1895. Fabaceae. = A. chamaeleuce Gray Garfield Co., Henry Mts., Jones 5658q, 1894 (POM!). Astragalus coltonii Jones var. aphyllus Jones Rev. As- tragalus 71. 1923. Fabaceae. Syn: A. coltonii var. coltonii Sevier Co., near Richfield, Jones sn, 1898 (POM!). Astragalus coltonii Jones var. coltonii Zoe 2: 237. 1891. Fabaceae. SynAstragalus coltonii var. aphyllus Jones Carbon Co., Castle Gate, Jones sn, 1889 (POM!;US!;NY!). Astragalus coltonii Jones var. moabensis Jones Contr. W. Bot. 8: 11. 1898. Fabaceae. Syn: Homalobus canovi- rens Rydb. San Juan Co., Monticello, Eastwood 9, 1892 (POM!). Astragalus convallarius Greene var. finitimus Barn- eby Leafl. W. Bot. 7: 192. 1954. Fabaceae. Washington Co., 3 mi s Enterprise, Ripley & Barneby 4967, 1942 (CAS!). Astragalus cottamii Welsh Rhodora 72: 189. 1970. Fa- baceae. San Juan Co., e Clay Hills Divide, Welsh 5207, 1966 (BRY!;NY!;ISC!). Astragalus cronquistii Barneby Mem. New York Bot. Card. 13: 258. 1964. Fabaceae. San Juan Co., w side Comb Wash, Cronquist 9123, 1961 (NY!;UTC!). Astragalus cymboides Jones Proc. Calif. Acad. II, 5: 650. 1895. Fabaceae. Syn: A. amphyoxys var. cymbellus Jones Emery Co., Huntington, Jones 5464j, 1894 (POM!;US!;N'y!). Astragalus desereticus Barneby Mem. New York Bot. Card. 13: 635. 1964. Fabaceae. Sanpete Co., Indianola, Tidestrom 2249, 1909 (GH). Astragalus desperatus Jones var. desperatus Zoe 2: 243. 1891. Fabaceae. Grand Co., near Cisco, Jones sn, 1890(US!;RM!;BRY!). Astragalus desperatus Jones var. petrophilus Jones Rev. Astragalus 204. 1923. Fabaceae. Emery Co., San Rafael Swell, Jones sn, 1914 (POM!;RM!;BRY!;NY!). Astragalus detritalis Jones Contr. W. Bot. 13: 9. 1910. Fabaceae. Syn: A. spectabilis C.L. Porter Duch- esne Co., near Theodore, Jones sn, 1908 (POM!;BRY!;NY!). Astragalus diehlii Jones Rev. Astragalus 194. 1923. Fabaceae. = A. flexuosus var. diehlii (Jones) Barneby Carbon Co., Farnham, Jones sn, 1898 (POM!;RM!; NY!;BRY!;UTC!). Astragalus dodgeanus Jones Zoe 3: 289. 1893. Fa- baceae. = A. wingatanus Wats. Grand Co., Thompsons Springs, Jones sn, 1891 (POM!;BRY!;NY!). Astragalus duchesnensis Jones Contr. W. Bot. 13: 6. 1910. Fabaceae. Duchesne Co., Theodore to Myton, Jones sn, 1908 (POM!;CAS!;RM!;BRY!;US!;NY!). Astragalus ensiformis Jones var. gracilior Barneby Proc. Calif. Acad. IV. 25: 158. 1944. Fabaceae. = A. en- siformis Jones Washington Co., s Veyo, Ripley & Barn- eby 4951, 1942 (CAS!;NY!). Astragalus episcopus Wats. Proc. Amer. Acad. 10: 345. 1875. Fabaceae. Kane (?) Co., Bishop sn, 1872 (GH;US!;NY!). Astragalus equisolensis Neese & Welsh Rhodora 83: 457. 1981. Fabaceae. Uintah Co, s Jensen, Neese & Welsh 7380, 1979 (BRY!;NY!;US!;POM!;UTC!). Astragalus eremiticus Sheld. Bull. Geol. & Nat. Hist. Surv. Minnesota 9: 161. 1894. Fabaceae. Washington Co., Beaver Dam Mountains, Parry 45, 1874 (MINN; US!;ISC!;NY!). Astragalus eurekensis Jones Contr. W. Bot. 8: 12. 1898. Fabaceae. Syn: Xylophacos medius Rydb. Juab Co., Eureka, Jones sn, 1891 (POM!;BRY!). Astragalus flavus Nutt. in T. & G. var. candicans Gray Proc. Amer. Acad. 12: 54. 1876. Fabaceae. Syn: A. confertiflorus Gray Sevier Co., near Richfield, Ward 246, 1875 (GH;NY!). Astragalus hamiltonii C.L. Porter Rhodora 54: 159. 1952. Fabaceae. Uintah Co., 5 mi sw Vernal, Hamilton & Beathsn, 1952 (RM!). Astragalus harrisonii Barneby Mem. New York Bot. Card. 13: 271. 1964. Fabaceae. Wayne Co., near Fruita, Barneby 13131, 1961 (CAS!;US!;BRY!;NY!;UTC!). Astragalus haydenianus Gray ex Brandegee var. ma- jor Jones Zoe 2: 240. 1891. Fabaceae. = A. bisulcatus var. major (Jones) Welsh Kane Co., Johnson, Jones sn, 1890(POM!;BRY!;NY!). Astragalus ibapensis Jones Zoe 3: 290. 1893. Fa- baceae. = A. diversifolius Gray Juab Co., Deep Creek Mts., Jones sn, 1891 (POM!). Astragalus iselyi Welsh Great Basin Naturalist 34: 305. 1974. Fabaceae. San Juan Co., Brumley Ridge, Welsh 10970, 1971 (BRY!;UT!;POM!;UTC!;ISC!). Astragalus jejunus Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 73. 1871. Fabaceae. Summit (?) Co., Bear River Valley, Watson 279, 1869 (NY!). Astragalus junceus var. attenuatus Jones Rev. Astra- galus 76. 1923. Fabaceae. = A. convallarius Greene Carbon Co., Price, Jones sn, 1898 (POM!;BRY!;NY!). Astragalus lentiginosus Dougl. ex Hook. var. char- taceus Jones Proc. Calif. Acad. II. 5: 673. 1895. Fa- baceae. = A. lentiginous var. araneosus (Sheld.) Barn- eby Sanpete Co., Ephriam, Jones 5627m, 1894 (POM!). Astragalus lentiginosus Dougl. ex Hook. var. pohlii Welsh & Barneby Isleya 2: 1. 1981. Fabaceae. Tooele Co., 4.5 mi N Vernon, Welsh et al. 16743, 1978 (BRY!;NY!;UTC!). June 1982 Welsh: Utah Plant Types 161 Astragalus lentiginosus Dougl. ex Hook. var. vitreus Bameby Leafl. W. Bot. 4: 119. 1945. Fabaceae. Wash- ington Co., 5 mi w Leeds, Maguire & Blood 4413, 1933 (UTC!). Astragalus lentiginosus Dougl. ex Hook. var. wah- weapensis Welsh Great Basin Naturahst 38: 286. 1978. Fabaceae. Kane Co., Four Mile Bench, Welsh 12426, 1974 (BRY!;ISC!). Astragalus limnocharis Bameby Leafl. W. Bot. 4: 236. 1946. Fabaceae. Kane Co., Navajo Lake, Maguire 19474, 1940(NY!;RM!;UTC!). Astragalus nidularius Bameby Leafl. W. Bot. 8: 16. 1956. Fabaceae. San Juan Co., White Canyon, Barneby 12778, 1955 (CAS!;US!;RM!;NY!;UT!;UTC!)'. Astragalus palans Jones Zoe 4: 37. 1893. Fabaceae. = A. lentiginosus var. palans (Jones) Jones San Juan Co., Montezuma Canyon, Eastwood sn, 1892 (RM!;NY!). Astragalus peabodianus Jones Zoe 3; 295. 1893. Fa- baceae. = A. pubentissinuis var. peabodianus (Jones) Welsh Grand Co., Thompsons Springs, Jones sn, 1891 (POM!). Astragalus perianus Barneby Mem. New York Bot. Card. 13(2): 973. 1964. Fabaceae. Piute Co, n Bullion Creek, Rydberg & Carlton 7104, 1905 (NY!;RM!;US!). Astragalus pictus var. angustus Jones Zoe 4: 37. 1893. Fabaceae. = A. ceramicus Sheld. San Juan Co., Mon- tezuma Canyon, Eastwood 7, 1892 (POM!;US!). Astragalus pictus var. magnus Jones Rev. Astragalus 109. 1923. Fabaceae. = A. ceramicus Sheld. Washington Co., Silver Reef, Jones 5160, 1894 (POM!;RM!). Astragalus pinonis Jones Contr. W. Bot. 8: 14. 1898. Fabaceae. Beaver Co., Frisco, Jones sn, 1880 (POM!). Astragalus preussii Gray var. latus Jones Zoe 4: 36. 1893. Fabaceae. = A. preussii Gray var. preussii Emery Co., Green River, Jones sn, 1891 (NY). Astragalus preussii Gray var. sulcatus Jones Zoe 4: 37. 1893. Fabaceae. = A. eastwoodiae ' ]ones Southeast Utah, Cane Spring, Eastwood sn, 1892 (CAS!;BRY!; NY!;POM!). Astragalus rafaelensis Jones Rev. Astragalus 146. 1923. Fabaceae. Emery Co., Cedar Mt., Jones sn, 1915 (POM!;CAS!;RM!;BRY!;NY!;UTC!). Astragalus sabulosus Jones Zoe 2: 239. 1891. Fa- baceae. Grand Co., Cisco, Jones sn, 1890 (POM!;BRY!). Astragalus saurinus Barneby Leafl. W. Bot. 8: 17. 1956. Fabaceae. Uintah Co, 6 mi N Jensen, Holmgren & Tillett 9527, 1953 (NY!;CAS!;RM!). Astragalus serpens Jones Proc. Calif. Acad. II, 5: 641. 1895. Fabaceae. Wayne Co., Loa Pass, Jones 56.39i, 1894 (POM!;US!;NY!). Astragalus sesquiflorus Wats. Proc. Amer. Acad. 10: 345. 1875. Fabaceae. Kane Co., near Kanab?, Bishop sn, 1873 (GH;US!). Astragalus sileranus Jones Zoe 2: 242. 1891. Fa- baceae. = A. subcinereus Gray Kane Co., Sink Valley, Jones sn, 1890 (POM!;CAS!;US!;BRY!;NY!). Astragalus sileranus Jones var. caraicus Jones Proc. Calif. Acad. II. 5: 642. 1895. Fabaceae. = A. sub- cinereus Gray Kane (?) Co., Elk Ranch, Jones 6036, 1894 (POM!;BRY!). Astragalus spectabilis C.L. Porter Rhodora 54; 160. 1952. Fabaceae. = A. detritahs Jones Uintah Co., 5 mi sw Vernal, Porter 5309, 1950 (RM!;CAS!;BRY!; POM!;NY!). Astragalus stocksii Welsh Great Basin Naturalist 34: 307. 1974. Fabaceae. = A. henriinontanensis Welsh Garfield Co., Henry Mts., Welsh 11740, 1972 (BRY!;ISC!). Astragalus straturensis Jones Contr. W. Bot. 8: 19. 1898. Fabaceae. Washington Co., Silver Reef, Jones 5175, 1894 (POM!;US!;RM!;NY!;BRY!). Astragalus striatiflorus Jones Proc. Calif. Acad. II. 7: 643. 1895. Fabaceae. Washington Co., Springdale, Jones 6080k, 1894 (POM). Astragalus subcinereus Gray var. basalticus Welsh Great Basin Naturalist 38: 302. 1978. Fabaceae. Sevier Co., 10 mi s Fremont Jet., Welsh et al. 6447, 1967 (BRY!;ISC!). Astragalus subcinereus Gray var. subcinereus Proc. Amer. Acad. 13: .366. 1878. Fabaceae. Syn: A. sileranus Jones; A. sileranus var. caraicus Jones Southern Utah, (Northern Arizona ?), Palmer 117 1877 (US!;ISC!;NY!). Astragalus tegetarius Wats. var. rotundus Jones Proc. Calif. Acad. II. 5; 650. 1895. Fabaceae. = A. kent- rophyta var. implexus (Canby) Barneby Wayne Co., near Loa, Jones 5649b, 1894 (POM!;UC!). Astragalus tetrapterus Gray Proc. Amer. Acad. 13: .369. 1878. Fabaceae. Washington Co., 25 mi n St. George, Palmer 111, 1877 (NY!;ISC!). Astragalus thompsonae Wats. Proc. Amer. Acad. 10; .344. 1875. Fabaceae. = A. moUissimus var. thompsonae (Wats.) Barnebv Kane Co., Kanab, Thompson sn, 1872 (GH;US!). Astragalus ursinus Gray Proc. Amer. Acad. 13: 367. 1878. Fabaceae. = A. lentiginosus var. ursinus (Gray) Barneby Iron (?) Co., Bear Valley, Palmer sn, 1877 (GH!;NY!;ISC!). Astragalus wardii Gray Proc. Amer. Acad. 12: 55. 1876. Fabaceae. Garfield (?) Co., Aquarius PI., Ward 424, 1875 (GH;US!;NY!). Astragalus woodruffii Jones Rev. Astragalus 77. 1923. Fabaceae. Emery Co., San Rafael Swell, Jones sn, 1914 (POM!;US!;CAS!';RM!;BRY!;NY!;UTC!). Astragalus zionis Jones Proc. Calif. Acad. II, 5: 652. 1895. Fabaceae. Washington Co., Springdale, Jones 5261w, 1894 (POM!;US!;RM!;BRY!). Atriplex bonnevillensis C.A. Hanson Stud. Syst. Bot. (BRY) 1: 2. 1962. Chenopodiaceae. Millard Co., Pine Valley playa, Hanson 354, 1960 (BRY!;UTC!). Atriplex caput-medusae Eastw. Proc. Calif. Acad. II. 6: 316. 1896. Chenopodiaceae. = A. argentea var. caput- medusae (Eastw.) Fosberg San Juan Co., Recapture Creek, Eastwood 116, 1895 (US!;CAS!). Atriplex cornuta Jones Proc. Calif. Acad. II. 5; 718. 1895. Chenopodiaceae. = A. saccaria Wats. Emery Co., Green River, Jones 5481, 1894 (POM!;US!). Atriplex cuneata A. Nels. Bot. Gaz. 34: 357. 1902. Chenopodiaceae. Emery Co., Emery, Jones 5443, 1894 (US!;RM!). Atriplex cuneata A. Nels. ssp. introgressa C.A. Han- son Stud. Syst. Bot. (BRY) 1: 4. 1962. Chenopodiaceae. Carbon Co., Wellington, Hanson 346, 1961 (BRY!;POM!). Atriplex garrettii Rydb. Bull. Torrey Bot. Club 39; 312. 1912. Chenopodiaceae. Grand Co., Moab vicinity, Rydberg & Garrett 8465, 1911 (NY!;US!;UT!). Atriplex graciliflora Jones Proc. Calif. Acad. II, 5: 717. 1895. Chenopodiaceae. Wayne Co., Blue Valley, Jones 5697, 1894 (POM!;US!;RM!;BRY!). 162 Great Basin Naturalist Vol. 42, No. 2 Atriplex nuttallii Wats. var. utahensis Jones Contr. W. Bot. 11: 19. 1903. Chenopodiaceae. = A. tridentata Kuntze Salt Lake Co., Salt Lake City, Jones 1760, 1879? (UC). Atriplex rydbergii Standi. N. Amer. Fl. 21: 47. 1916. Chenopodiaceae. = A. argentea var. argentea San Juan (?) Co., s Moab, Rydberg & Garrett 9110, 1911 (US!;NY!). Atriplex subdecumbens Jones Proe. Calif. Acad. II. 5: 716. 1895. Chenopodiaceae. = A. truncata (Torr.) Gray Sevier Co., Fish Lake, Jones 5745, 1894 (POM!;US!). Atriplex tenuissima A. Nels. Bot. Gaz. 34: 359. 1902. Chenopodiaceae. = A. wolfii Wats. Sanpete Co., Gun- nison, Jones 6.525, 1900 (US!;RM!;POM!;NY!). Atriplex tridentata Kuntze Rev. Gen. 2: 546. 1891. Chenopodiaceae. Syn: A. nuttallii var. utahensis Jones Box Elder Co., Corinne, Kuntze .3084, 1874 (NY!). Atriplex welshii C. A. Hanson Stud. Syst. Bot. (BRY) 1: 1. 1962. Chenopodiaceae. Grand Co., 4 mi s Cisco, Hanson ,322, 1961 (BRY!;ISC!). Atropis laevis var. rigida Beal Grasses N. Amer. 2: 578. 1896. Poaceae. = Poa canbyi (Scribn.) Howell Tooele Co., Lake Point, Jones 1021, 1879(US!). Aulospermum minimum Mathias Ann. Missouri Bot. Card. 17: 353. 1930. Apiaceae. = Cijmopterus minimus (Mathias) Mathias Iron Co., Cedar Breaks, Mathias 723, 1929 (MO;CAS!;BRY!). Aulospermum rosei Jones in Coult. & Rose Contr. U.S. Natl. Herb. 7: 179. 1900. Apiaceae. = Cijmopterus rosei (Jones) Jones Sevier Co., Richfield, Jones 30, 1899 (US!;POM!). Bahia desertorum Jones Zoe 2: 249. 1891. Asteraceae. = Platyschkuhria integrifolia var. deserertorum (Jones) Ellison Grand Co., Cisco, Jones sn, 1890 (US!;POM!). Bahia ourolepis Blake Proc. Biol. Soc. Washington 35: 175. 1922. Asteraceae. = Platyschkuhria oblongifolia (Gray) Rydb. Emery Co., Green River, Jones 5482#1 1894 (US!;POM!). Bahamorhiza hispidula Sharp Ann. Missouri Bot. Card. 22: 137. 1935. Asteraceae. = B. hookeri var. hispi- dula (Sharp) Cronq. Tooele Co., Lake Point, Jones 1727, 1880 (UT!;UTC!). Batidophaca sabinarum Rydb. N. Amer. Fl. 24: 320. 1929. Fabaceae. = Astragalus argophyllus var. pan- guicensis (Jones) Jones Iron Co., Cedar Canyon, Garrett 2660, 1920 (NY!). Betula utahensis Britt. Bull. Torrey Bot. Club 31: 165. 1904. Betulaceae. = B. occidentalis x B. papyrifera Salt Lake Co., City Creek Canyon, Stokes sn, 1900 (NY!). Bigelovia douglasii var. spathulata Jones Proc. Calif. Acad. II, 5: 690. 1895. Asteraceae. = Chrysothamnus viscidiflorus var. pumilus (Nutt.) Jeps. Sevier Co., Fish Lake, Jones 5758m, 1894 (POM;US!). Bigelovia glareosa Jones Zoe 2: 247. 1891. Asteraceae. = C. nauseosus (Pallas) Britt. var. glareosa (Jones) Welsh Piute Co., Marysvale, Jones sn, 1890 (POM). Bigelovia howardii var. attenuata Jones Proc. Calif. Acad. II, 5: 691. 1895. Asteraceae. = Chrysothamnus parryi var. attenuatus (Jones) Kittell Piute Co., Mary- svale, Jones 5912, 1894 (US!;RM!;NY!). Bigelovia leiosperma Gray Syn. Fl. 1(2): 129. 1884. As- teraceae. = Chrysothamnus nauseosus var. leiosperma (Gray) Hall Washingto n Co., St. George, Palmer sn, 1875 (NY!;BRY!). Bigelovia leiosperma Gray var. abbreviata Jones Proc. Calif. Acad. II, 5: 693. 1895. Asteraceae. = Chrysothamnus nauseosus (Pallas) Britt. Sevier Co., Clear Creek Canyon, Jones 6105, 1894 (POM!;US!;NY!). Bigelovia menziesii var. scopuhrum Jones Proc. Calif. Acad. II, 5: 692. 1895. Asteraceae. = Haplopappus sco- pulorum (Jones) Blake Iron Co., near Cedar City, Jones 5204v, 1894 (POM!;US!). Bigelovia turbinata Jones Proc. Calif. Acad. II, 5: 691. 1895. Asteraceae. = Chrysothamnus nauseosus var. turbinatus (Jones) Blake Garfield (?) Co., Canaan Ranch, Jones 6066c, 1894 (POM!;US!). BrickelUa atractyloides Gray Proc. Amer. Acad. 8: 290. 1870. Asteraceae. Utah?, Colorado River, Palmer sn, 1870 (US!). BrickelUa linifolia D.C. Eaton Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot. 5: 137. 1871. Asteraceae. = B. oblongifolia Wats. var. linifolia (D.C. Eaton) Robins. Utah Co., Jordan Valley, American Fork, Watson 493, 1869 (US!;CAS!). BrickelUa watsonii Robins. Mem. Gray Herb. 1: 42. 1917. Asteraceae. = B. microphylla var. watsonii (Rob- ins.) Welsh Utah Co., American Fork Canyon, Watson 494, 1869 (US!;NY!). Buddleja utahensis Gov. Proc. Biol. Soc. Washington 7: 69. 1892. Loganiaceae. Washington Co., St. George, Palmer 400, 1877 (US!;NY!;BRY!;ISC!). Caesalpinia repens Eastw. Zoe 4: 116. 1893. Fa- baceae. = Hoffmanseggia repens (Eastw.) Cockerell Grand Co., Courthouse Wash, Eastwood sn, 1892 (CAS!;US!). Calamagrostis scopulorum Jones Proc. Calif. Acad. II, 5: 722. 1895. Poaceae. Syn; C. scopulorum var. lucidu- lum Kearney Washington Co., Springdale, Jones 6075, 1894(POM!;NY!). Calamagrostis scopulorum Jones var. lucidula Kearney USD A Agrostol. Bull. 11: 33. 1898. Poaceae. = C. scopulorum Jones Salt Lake Co., Alta, Jones 1145, 1879 (POM!;NY!;BRY!;UTC!). Calochortus aureus Wats. Amer. Naturalist 7: 303. 1873. Liliaceae. Syn: C. nuttallii var. aureus (Wats.) Ownbey Kane (?) Co., "S. Utah" Thompson, 1872 (US!). Calochortus flexuosus Wats. Amer. Naturalist 7: 303. 1873. Liliaceae. Kane (?) Co., "S. Utah or N. Arizona", Thompson sn, 1872 (GH). Camassia quamash (Pursh) Greene ssp. utahensis Gould Amer. Midi. Naturalist 28: 740. 1942. Liliaceae. = C. quamash (Pursh) Greene Cache Co., Blacksmith Canyon, Maguire & Maguire 3265, 1932 (?). Camissonia gouldii Raven Contr. U.S. Natl. Herb. 37: 368. 1969. Onagraceae. Washington Co., N of St. George, Gould 1423, 1941 (US!;CAS!). Capnodes brachycarpum Rydb. Bull. Torrey Bot. Club 34: 426. 1907. Fumariaceae. = Corydalis caseana ssp. brachycarpa (Rydb.) M. Ownbey Salt Lake Co., Alta, Jones 1197, 1879 (NY!;US!;RM!;POM!;BRY!;UTC!). Cardamine cordifoUa Gray var. pubescens Gray ex Schuiz Engl. Bot. Jahrb. 32: 439. 1903. Brassicaceae. = C. cordifoUa Gray Garfield (?) Co., Thousand Lake Mtn., Ward .396, 1875 (US!). Cardamine palustris var. jonesii Kuntze Rev. Gen. 1: 125. 1891. Brassicaceae. = Rorippa curvipes var. cur- vipes Salt Lake Co., City Creek Canyon, Jones 1352, 1879(POM!;NY!). June 1982 Welsh: Utah Plant Types 163 Cardamine uinlahensis F.J. Hermann Rhodora 36: 410. 1934. Brassicaceae. = C. cordifolia Gray Summit Co., Mt. Elizabeth Ridge, Hermann 5894, 1933 (GH). Carduus lacerus Rydb. Bull. Torrey Bot. Club 37: 543. 1910. .A.steraceae. = Cirsium scariosum Nutt. Wasatch Co., near Midway, Carlton & Garrett 6732, 1905(RM!;NY!). Carduus olivescens Rydb. Bull. Torrey Bot. Club 37: 544. 1910. .\steraceae. = C. scariosum Nutt. Garfield Co., .\quarius Plateau, Rydberg & Carlton 7450, 1905 (NY!). €arex camptjlocarpa T.H. Holm ssp. affinis Maguire & Holmgren Leafl. W. Bot. 4: 262. 1946. Cyperaceae. = C. scopulorum T.H. Holm Juab Co., Indian Farm Creek, Maguire & Holmgren 21947. 1943 (NY!;US!;CAS!;UTC!). Carex canescens L. var. dubia Bailey Bot. Gaz. 9: 119. 1884. Cvperaceae. = C. cmiesceiis L. Summit (?) Co., Bear River Canyon, Watson 1231A, 1869 (NY!). Carex epapillosa Mack, in Rydb. Fl. Rocky Mts. 138. 1060. 1917. Cvperaceae. = C. atrata L. Piute Co., Mary- svale, Jones 5.345, 1894 (NY!;US!;POM!;BRY!). Carex interimus Maguire Brittonia 5: 200. 1944. Cvperaceae. = C. aquatilis Wahl. Cache Co, Tony Grove Lake, Maguire 16098, 19.38 (NY!;C.\S!;US!;UTC!)'. Carex pelocarpa F. Hermann Rhodora 39: 492. 1937. Cyperaceae. = C. nova Bailey Summit Co., Lamotte Peak, Hermann 5983, 19,33 (NY!';C.\S!). Carex rachillis Maguire Brittonia 5: 199. 1944. Cype- raceae. = C. sidmigricans Stacev Summit Co, Gilbert Peak. Maguire et al 14668, 1936 (NY!;US!;CAS!;UTC!). Carex vernacula Bailey var. hobsonii Maguire Brit- tonia 5: 199. 1944. Cvperaceae. = C. neurophora Mack. Cache Co., Bear River Range, Maguire et al 14013, 1936 (NY!;C.\S!;US!;UTC!). Carum garrettii A. Nels. in Coult. & Rose Contr. U.S. Natl. Herb. 12: 443. 1909. Apiaceae. = Perideridia gairdneri (H. & A.) Mathias Salt Lake (?) Co., Wasatch Mts.. Garrett 2053, 1906 (US!;NY!). Castilleja aquariensis N. Holmgren Bull. Torrey Bot. Club 100: 87. 1973. Scrophulariaceae. Garfield Co., Aquarius Plateau, N. & P. Holmgren 4726, 1970 (NY!;US!;BRY!;UTC!). Castilleja arcuata Rydb. Bull. Torrey Bot. Club 34: 35. 1907. Scrophulariaceae. = C. linariifolia Benth. Se- vier Co., Fish Lake, Rydberg & Carlton 7-508, 1905 (NY!:RM!:US!). Castilleja flava Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 2.30. 1871. Scrophulariaceae. Summit (?) Co., Upper Bear River Valley, Watson 813, 1869 (US!;NY!). Castilleja konardii Rydb. Bull. Torrey Bot. Club 34: .36. 1907. Scrophulariaceae. Utah Co., American Fork Canyon, Leonard 151, 1885 (NY!). Castilleja parvula Rydb. Bull. Torrey Bot. Club 34: 40. 1907. Scrophulariaceae. Piute Co., Bullion Creek, Rydberg & Carlton 71.58, 1905 (NY!;US!). Castilleja revealii N. Holmgren Bull. Torrey Bot. Club 100: 87. 1973. Scrophulariaceae. Garfield Co., Bryce Canyon, Holmgren & Reveal 2017, 1965 (NY!;US!;BRY!;UT!:UTC'). Castilleja variabilis Rydb. Bull. Torrey Bot. Club 34: ..37. 1907. Scrophulariaceae. = C. miniata Dougl. Salt Lake Co., Big Cottonwood Canyon, Rydberg 6773, 1905 (NY!;RM!). Castilleja viscida Rydb. Bull. Torrey Bot. Club 34: 32. 1907. Scrophulariaceae. Syn: C. applegatei var. vis- cida (Rydb.) Ownbey Salt Lake Co., Big Cottonwood Canyon, Rydberg & Carlton 6593, 1905 (NY!). Castilleja zionis Eastw. Leafl. W. Bot. 3: 91. 1941. Scrophulariaceae. Washington Co., Clear Creek, East- wood & Howell 9227, 1941 (CAS!). Caulanthus crassicaulis (Torr.) Wats. var. glaber Jones Zoe 4: 266. 1893. Brassicaceae. Kane Co., Sink Valley, Jones sn, 1890 (POM!). Caulanthus crassicaulis (Torr.) Wats. var. major Jones Proc. Calif. Acad. II, 5: 623. 1895. Brassicaceae. Garfield Co., Henry Mts., Jones 5685, 1894 (POM!;US!;BRY!;NY!). ' Caulanthus divaricatus Rollins Contr. Gray Herb. II, 201: 8. 1971. Brassicaceae. = Thelypodiopsis divaricata (Rollins) Welsh & Reveal San Juan Co., 10 mi E of Hite, Cronquist 90.33, 1961 (GH;NY!). Caulanthus hastatus Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 28. 1871. Brassicaceae. = Chloro- crambe hastatus (Wats.) Rydb. Salt Lake (?) Co., Wasatch Mts., Watson 114, 1869 (US!;NY!). Ceanothus fendleri Gray var. viridis Jones Proc. Ca- lif. Acad. II, 5: 629. 1895. Rhamnaceae. = C. fendleri Gray Kane (?) Co., Elk Ranch, Jones 60.32n, 1894 (POM!). Ceanothus martinii Jones Contr. W. Bot. 8: 41. 1898. Rhamnaceae. Syn: C. utahensis Eastw. Sanpete Co., Manti Canyon', Jones sn, 1895 (POM!;RM!;BRY!; NY!;UTC!). ' Ceanothus utahensis Eastw. Proc. Calif. Acad. IV. 16: 363. 1927. Rhamnaceae. = C. martinii Jones Wasatch Co., Soldier Summit, Eastwood 7689, 1918 (CAS!). Celtis villosula Rydb. Fl. Rocky Mts. ed 2. 1116. 1922. Ulmaceae. = C. reticulata Torr. Utah, without definite locality, Jones sn, 1894 (NY!). Cerastium variable Goodding Bot. Gaz. 37: 54. 1904. Carvophvllaceae. = C. beeringianum C. & S. Uintah Co.; Dyer Mine, Goodding 1246, 1902 (US!;RM!; BRY!;NY!;UT!;ISC!). Cercis orbiculata Greene Feddes Repert. 11: 111. 1912. Fabaceae. = C. occidentalis var. orbicidata (Greene) Tidestr. Washington Co., Diamond Valley, Goodding 899, 1902 (US!;NY!). Cercocarpus flabellifolius Rydb. N. Amer. Fl. 22: 422. 1913. Rosaceae. = C. montanus Raf. Sevier Co., near Glenwood, Ward 122, 1875 (US!;NY!). Cercocarpus intricatus Wats. Proc. .\mer. .\cad. 10: 346. 1875. Rosaceae. Syn: C. intricatus var. villosus Schneid. Utah Co., American Fork Canyon, Watson 314, 1869 (US!;NY!). Cercocarpus intricatus Wats. var. villosus Schneid. Mitt. Deutsch. Dendr. Ges. 14: 129. 1905. Rosaceae. = C. intricatus Wats. Tooele (?) Co., Deep Creek, Jones sn, 1891 (US!). Cercocarpus ledifolius Nutt. var. intercedens Schneid. f. hirsutus Schneid. Mitt. Deutsch. Dendrol. Ges. 14: 129. 1905. Rosaceae. = C. ledifolius Nutt. Weber Co., Ogden Canyon, Pammel & Blackwood 3726, 1902 (ISC!;BRY!). ' Cercocarpus ledifolius Nutt. var. intercedens f. sub- glaber Schneid. Mitt. Deutsch. Dendr. Ges. 14: 128. 1905. Rosaceae. = C. ledifolius Nutt. Utah Co., Slate Canyon, Jones 5613b, 1894 (MO;NYl;US!;RM!; BRY!;POM!). 164 Great Basin Naturalist Vol. 42, No. 2 Cercocarpus parvifolius Wool. var. minimus Schneid. 111. Handb. Laubholzk. 1: 532. 1905. Rosaceae. = C. montuntis Raf.? Utah ?. Chaenactis brachiata Greene Leafl. Bot. Obs. & Crit. 2: 224. 1912. Asteraceae. = C. douglasii (Hook.) H. & A. Washington Co., Springdale, Jones 5261q, 1894 (US!;POM!). Chaenactis brachiata Greene var. stansburiana Stockwell Contr. Dudley Herb. .3: 111. 1940. Asteraceae. = C. douglasii (Hook.) H. & A. Tooele Co., Stansbury Island, Stansbury sn, 1850 (US;NY!). Chaenactis douglasii (Hook.) H. & A. var. alpina Gray Syn. Fl. N. Amer. 1(2): 341. 1884. = C. alpina (Gray) Jones Salt Lake Co., Alta, Jones 1232, 1879 (NY!;UTC!). Cheirinia brachycarpa Rydb. Bull. Torrey Bot. Club 39: .325. 1912. Brassicaceae. = Erijsimum asperum Nutt. San Juan Co., Abajo Mts., Rydberg & Garrett 9713, 1911 (NY!;US!;UT!). Chrysopsis caespitosa Jones, not Nutt. Proc. Calif. Acad. II, 5: 694. 1895. Asteraceae. = Heterotheca pnesii (Blake) Welsh & Atwood Washington Co., Springdale, Jones .5249u, 1894 (POM!;US!). Chrysopsis villosa Nutt. var. cinerascens Blake Proc. Biol. Soc. Washington .35: 173. 1922. Asteraceae. = Het- erotheca villosa (Nutt.) Shinners Beaver Co., Beaver Canyon, Tidestrom 2873, 1901 (US!). Chrysopsis villosa Nutt. var. scabra Eastw. Proc. Ca- lif. Acad. II, 6: 294. 1896. Asteraceae. = Heterotheca vil- losa (Nutt.) Shinners San Juan Co., Willow Creek, East- wood .38, 1895 (CAS!). Chrysothamnus marianus Rydb. Bull. Torrey Bot. Club 37; 131. 1910. Asteraceae. = C. viscidiflorus var. puhendus (D.C. Eaton) Jeps. Piute Co., near Marysvale, Rydberg & Carlton 6993, 1905 (US;R]V1!;NY!). Chrysothamnus nauseosus (Pallas) Britt. ssp. iridis L.C. Anderson Great Basin Naturalist 41: 311. 1981. As- teraceae. Sevier Co., Rainbow Hills, Welsh 19258, 1979 (BRY!;FSU!). Chrysothamnus nauseosus (Pallas) Britt. var. psilo- carpus Blake J. Washington Acad. Sci. 27: 376. 1937. As- teraceae. Syn: C. nauseosus ssp. psilocarpus (Blake) L.C. Anderson Emery Co., Huntington Canyon, Garrett 7021 1935 (US). Chrysothamnus oliganthus A. Nels. Univ. Wyoming Publ. Bot. 1: 65. 1924. Asteraceae. = C. nauseosus var. leiospermus (Gray) Hall Washington Co., Zion National Park, Nelson 9975, 1922 (RM!). Chrysothamnus salicifolius Rydb. Bull. Torrey Bot. Club 37: 1,30. 1910. Asteraceae. = C. nauseosus (Pallas) Britt. var. salicifolius (Rydb.) Hall Wasatch Co., Straw- berry Valley, Leonard 288, 1883 (NY!;BRY!). Chrysothamnus zionis A. Nels. Univ. Wyoming Publ. Bot. 1: 66. 1924. Asteraceae. = C. nauseosus var. gnaphaloides (Greene) Hall Washington Co., 20 mi N St. George, Nelson 9980, 1922 (RM!). Cirsium barnebyi Welsh & Neese in Welsh Brittonia 33: 296. 1981. Asteraceae. Uintah Co., 1.5 mi e Ignacio, Welsh 19606, 1980 (BRY!;NY!;US!;MINN!). Cirsium eHocephalum var. leiocephalum D.C. Eaton Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 196. 1871. Asteraceae. = C. eatonii Robins. var. eatonii Summit? Co., Uinta Mts., Watson 691, 1869 (US!;NY!). Cirsium lactucinum Rydb. Fl. Rocky Mts. 1010, 1068. 1917. Asteraceae. = C. rijdbergii Petrak San Juan Co., Bluff, Rydberg & Garrett 10001, 1911 (NY!;US!;UT!). Cirsium pulchellum (Greene) Woot. & Standi, var. glabrescens Petrak Beih. Bot. Centr. 3.5(2): 511. 1917. A.steraceae. = C. bipinnatum (Eastw.) Petrak San Juan Co., Elk Mts., Rydberg & Garrett 9335, 1911 (US!). Cirsium rydbergii Petrak Beih. Bot. Centralbl. 35(2): 315. 1917. Asteraceae. Syn: C. lactucinum Rydb. San Juan Co., Bluff, Rydberg & Garrett 10001, 1911 (US!;UT!;NY!). Cirsium undulatum (Nutt.) Spreng. var. albescens D.C. Eaton Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 194. 1871. Asteraceae. = C. undulatum (Nutt.) Spreng. Tooele Co., Stansbury Island, Wat.son 687, 1869 (US!). Cirsium utahense Petrak Beih. Bot. Centr. 35(2); 470. 1917. Asteraceae. Washington Co., Silver Reef, Jones 516.3q, 1894 (POM !;US!). Clematis alpina ssp. occidentalis var. repens Kuntze Verb. Bot. Ver. Brandenb. 26: 161. 1885. Ranuncu- laceae. = C. columbiana (Nutt.) T. & G. Utah Co., American Fork, Jones 1887, 1880 (NY!;POM!). Clematis douglassii ssp. jonesii Kuntze Verb. Bot. Ver. Brandenb. 25: 180. 1886. Ranunculaceae. = C. hir- sutissima Pursh Utah Co., American Fork Canvon, Jones 1351, 1880 (US!;POM!;NY!). Clematis pseudotragene ssp. tvenderothioides Kuntze Verb. Bot. Ver. Brandenb. 26: 160. 1885. Ranuncu- laceae. = C. columbiana var. columbiana Kane Co., Ka- nab, Siler sn, 1873 (K). Cleome integrifolia var. angusta Jones Proc. Calif. Acad. II, 5; 625. 1895. Capparidaceae. = C. serrulata var. angusta (Jones) Tidestr. Piute Co., Marysvale, Jones 6057, 1894 (US!). Cleomella cornuta Rydb. Bull. Torrey Bot. Club .30; 249. 1903. Capparidaceae. = C. palmeriana Jones Wayne Co., Caineville, Jones 5656, 1894 (US!;BRY!;RM!;NY!). Cleomella nana Eastw. Bull. Torrey Bot. Club 30: 490. 1903. Capparidaceae. = C. palmeriana Jones Grand Co., S of Thompsons Spr., Eastwood sn, 1892 (CAS!;RM!). Cleomella palmerana Jones Zoe 2: 236. 1891. Cap- paridaceae. Emery Co., Green River, Jones sn, 1890 (POM;US!). Cnicus calcareus Jones Proc. Calif. Acad. II, 5: 704. 1895. Asteraceae. = Cirsium calcareum (Jones) Woot. & Standi. Garfield Co., Bromide Pass, Jones 569.5bh, 1894 (POM!;BRY!;US!;NY!). Cnicus clavatus Jones Proc. Calif Acad. II, 5; 704. 1895. Asteraceae. = Cirsium clavatum (Jones) Petrak Se- vier Co., Fish Lake, Jones 5715, 1894 (POM!;BRY!;US!;NY!). Cnicus nidulus Jones Proc Calif. Acad. II, 5: 705. 1895. Asteraceae. = Cirsium nidulum (Jones) Petrak Garfield Co., Paria, Jones 5290a, 1894 (POM!;US!;NY!). Cnicus rothrockii var. diffusus Eastw. Proc. Calif. Acad. II, 6; 303. 1896. Asteraceae. = C. rothrockii (Gray) Petrak San Juan Co., Willow Creek, Eastwood 65, 1895 (US!;CAS!). Cogswellia cottamii Jones Contr. W. Bot. 16: .36. 1930. Apiaceae. = Lomatium parriji (Wats.) Macbr. Washington Co., Beaverdam Mts., Cottam et al. 4098, 1929 (BRY!;UT!). June 1982 Welsh: Utah Plant Types 165 Cogswellia millefolia var. depauperata Jones Contr. W. Bot. 12: 38. 1908. Apiaceae. = Lomatium graiji Coiilt. & Rose Tooele Co., Dugway, Jones sn, 1891 (US!;POM!;NY!). Cogswellia minima Mathias Ann. Missouri Bot. Card. 19: 497. 1932. Apiaceae. = Lomatium rninimtan (Math- ias) Mathias Garfield Co., Bryce Canyon, Mathias 670, 1929 (MO;CAS!;NY!). Coleosanthes garretti A. Nels. Proc. Biol. Soc. Wash- ington 20: 38. 1907. Asteraceae. = Brickellia grandiflora (Hook.) Niitt. Salt Lake Co., City Creek Canyon, Gar- rett 1061, 1904 (RM!;UT!). Collomia tenella Gray Proc. Amer. Acad. 8: 259. 1870. Polemoniaceae. Summit Co., Parleys Park, Watson 900, 1869 (NY!). Coloptera jonesii Coult. & Rose Rev. N. Amer. Um- bell. 49. 1888. Apiaceae. = Cymopterus newberriji (Wats.) Jones Beaver Co., Milford, Jones 1792, 1880 (POM!;UT!). Comandra linearis Rydb. Fl. Rocky Mts. 818. 1066. 1917. Santalaceae. = C. uynbellata var. pallida (DC.) Jones Emery Co., Green River, Tracy 716, 1887 (NY!). Cordylanthus parryi Wats, in Parry Amer. Naturalist 9; 346. 1875. Scrophulariaceae. = C. maritimus ssp. ca- nescens (Gray) Chuang & Heckard Washington Co., Parry 155, 1874 (US!;NY!;ISC!). Corydalis engelmannii var. exaltata Fedde Feddes Repert. 11: 497. 1913. Fumariaceae. = C. aurea Willd. Grand (?) Co., La Sal Mts., Purpus 6550, 1897 (US!). Coryphantha marstonii Clover Bull. Torrey Bot. Club 65: 412. 1938. Cactaceae. = C. Tuissouriensis var. marstonii (Clover) L. Benson Garfield Co., Hells Back- bone, Clover 1909, 1937 (?) (MICH). Cowania stansburiana Torr. in Stansb. Explor. Great Salt Lake 386. 1852. Rosaceae. = C. mexicana D. Don Davis (?) Co., Great Salt Lake, Stansbury sn, 1850 (NY!). Crassipes annuus Swallen Amer. J. Bot. 18: 685. 1931. Poaceae. = Sclerochloa dura (L.) Beauv. Davis (?) Co., between Salt Lake City & Ogden, Fallas sn, 1928 (US!). Crepis aculeolata Greene Leafl. Bot. Obs. & Grit. 2: 86. 1910. Asteraceae. = C. runcinata var. runcinata Utah, Ward 606, 1875 (US!). Crepis occidentalis Nutt. var. costatus Gray Geol. Surv. Calif. Bot. 1: 435. 1880. Asteraceae. = C. occiden- talis Nutt. Tooele Co., Stansbury Island, Watson 715, 1869 (US!). Cressa erecta Rydb. Bull. Torrey Bot. Club 40: 466. 1913. Convolvulaceae. = C. truxillensis H.B.K. Salt Lake Co., Becks Hot Springs, Garrett 870f, 1905 (NY!). Croton longipes Jones Proc. Calif. Acad. II, 5: 721. 1895. Euphorbiacea. Washington Co., 2 mi e Leeds, Jones 5213, 1894 (POM!;US!;CAS!;RM!;NY!). Cryptantha bamebyi Johnst. J. Arnold Arb. 29: 240. 1948. Boraginaceae. Uintah Co., 30 mi s Ouray, Ripley & Barneby 8748, 1947 (GH;NY!). Cryptantha compacta Higgins Great Basin Naturalist 28: 196. 1968. Boraginaceae. Millard Co., 8 mi w Desert Experimental Range, Higgins 1613, 1963 (BRY!;US!;POM!). Cryptantha grahamii Johnst. J. Arnold Arb. 18: 23. 1937. Boraginaceae. Uintah Co., mouth Sand Wash, Graham 7924, 1933 (GH). Cryptantha johnstonii Higgins Great Basin Naturalist 28: 195. 1968. Boraginaceae. Emery Co., 15 mi w Hwy 50-6, Higgins 1310,1968 (BRY!;POM!;US!;NY!). Cryptantha leptophylla Rydb. Bull. Torrey Bot. Club 36: 679. 1909. Boraginaceae. = C. nevadensis Nels. & Kennedy Washington Co., St. George, Palmer 350, 1877 (NY!). Cryptantha ochroleuca Higgins Great Basin Natural- ist 28: 197. 1968. Boraginaceae. Garfield Co., Red Can- yon, Higgins 1788, 1968 (BRY!;US!). Cryptantha rollinsii Johnst. J. Arnold Arb. 20: 391. 1939. Boraginaceae. Uintah Co., 22 mi s Ouray, Rollins 1715, 1937 (GH;US!;CAS!;RM!;UTC!). Cuscuta denticulata Engelm. in Parry Amer. Natural- ist 9: 348. 1875. Cuscutaceae. Washington Co., near St. George, Parry 205, 1874 (US!;NY!;ISC!;BRY!). Cuscuta warneri Yuncker Brittonia 12: 38. 1960. Cus- cutaceae. Millard (?) Co., Flowell, Warner sn, 1957 (US!;CAS!;BRY!;UT!;NY!;UTC!;ISC!). Cycladenia jonesii Eastw. Leafl. W. Bot. 3: 159. 1942. Apocynaceae. = C. humilis var. jonesii (Eastw.) Welsh & Atwood Emery Co., San Rafael Swell, Jones sn, 1914 (CAS!;US!;BRY!). Cymopterus basalticus Jones Contr W. Bot. 12: 16. 1908. Apiaceae. Beaver (?) Co., Wah Wah, Jones sn, 1906 (POM !;BRY!). Cymopterus beckii Welsh & Goodrich in Welsh Brit- tonia 33: 297. 1981. Apiaceae. Wayne Co., Fruita, Beck sn, 1938 (BRY!). Cymopterus corrugatus var. scopulicola Jones Contr. W. Bot. 14: 39. 1912. Apiaceae. = C. coulteri (Jones) Mathias Juab Co., Sevier Bridge, Jones sn, 1910 (POM!). Cymopterus decipiens Jones Zoe 2: 247. 1891. Apiaceae. = C. fendleri Gray Grand Co., Cisco, Jones sn, 1891 (POM!;US!;NY!). Cymopterus duchesnensis Jones Contr. W. Bot. 13: 12. 1910. Apiaceae. Duchesne Co., Myton, Jones sn, 1908 (CAS!;US!;RM!;POM!;BRY!). Cyrhopterus higginsii Welsh Great Basin Naturalist 35: 377. 1976. Apiaceae. Kane Co., 17 mi e Glen Canyon City, Welsh 12740, 1975 (BRY!;NY!). Cymopterus ibapensis Jones Zoe 3: 302. 1893. Apiaceae. Tooele (?) Co., Deep Creek Valley, Jones sn, 1891 (POM!;US!;RM!). Cymopterus jonesii Coult. & Rose Rev. N. Amer. Um- bell. 80. 1888. Apiaceae. Syn: Aulospermum jonesii (Coult. & Rose) Coult. & Rose Beaver Co., Frisco, Jones 1808, 1880 (US!;CAS!;POM!;RM!;NY!;UT!;UTC!). Cymopterus longipes Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 124. 1871. Apiaceae. Syn: Peuceda- num lapidosum Jones S.alt Lake Co., near Salt Lake City, Watson 451, 1869 (US!;NY!). Cynomarathrum latilobum Rydb. Bull. Torrey Bot. Club 40: 73. 1913. Apiaceae. = Lomatium latilobum (Rydb.) Mathias Grand Co., Wilson Mesa, Rydberg & Garrett 8371, 1911 (NY!;US!;UT!). Cynomarathrum scabrum Coult. & Rose Contr. U.S. Natl. Herb. 7: 247. 1900. Apiaceae. = Lomatium sca- brum (Coult. & Rose) Math. & Const. Beaver Co., Frisco, Jones 1864, 1880 (US!;CAS!;POM!;BRY!; NY!;UT!;UTC!). Cystium stramineum Rydb. N. Amer. Fl. 24: 409. 1929. Fabaceae. = Astragalus lentiginosus var, stra- mineus (Rydb.) Barneby Washington Co., "S. Utah" Pal- mer sn, 1870 (NY!;US!). Dalea epica Welsh Great Basin Naturalist 31: 90. 1971. Fabaceae. San Juan Co., 10 mi e Halls Crossing, Welsh 5205, 1966 (BRY!;NY!;ISC!). 166 Great Basin Naturalist Vol. 42, No. 2 Dalea johnsonii Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 64. 1871. Fabaceae. = Psorotharnnus fremontii (Torr.) Barneby Washington Co., near St. George, Johnson sn, 1870 (GH). Dalea nummularia Jones Contr. W. Bot. 18: 41. 1933. Fabaceae. = Psorotharnnus polyadenius var. pnesii Barneby Emery Co., Green River, Jones sn, 1914 (US!;CAS!;NY!).' Daucophyllum lineare Rydb. Bull. Torrey Bot. Club 40: 69. 1913. Apiaceae. = Musineon lineare (Rydb.) Mathias Cache Co., near Logan, Rydberg sn, 1895 (?). Delphinium ahietorum Tidestr. Proc. Biol. Soc. Washington 27: 61. 1914. Ranunculaceae. = D. occiden- tale Wats. Sanpete (?) Co., Wasatch Plateau, Coville & Tidestrom 19, 1908 (US!). Delphinium coelestinum Rydb. Bull. Torrey Bot. Club .39: 320. 1912. Ranunculaceae. Syn: D. ajnabile Tidestr. (new name) = D. scaposum Greene Washing- ton Co., St. George, Palmer 10, 1877 (NY!;US!). Delphinium leonardii Rydb. Bull. Torrey Bot. Club 39: .320. 1912 Ranunculaceae. = D. andersonii Gray Salt Lake Co., Garfield, Leonard 20.5, 1884 (NY!). Delphituum scoptilorum var. attenuafum Jones Proc. Calif. Acad. II, 5: 617. 1895. Ranunculaceae. = D. bor- heifi Huth. Piute Co., head Bullion Creek, Jones 5893d, 1894(POM!:US!). Descurainia richardsonii (Sweet) Schulz var. macros- perma Schulz Pflanzenr. 4. Fam. 105(Heft 86): 319. 1924. Brassicaceae. = D. richardsonii var. brevipes (Nutt.) Welsh & Reveal Salt Lake Co., Alta, Jones 1117, 1879 (UTC!). Dicoria paniculata Eastw. Proc. Calif. Acad. II, 6: 298. 1896. Asteraceae. = D. brandegei Gray San Juan Co., McElmo & Recapture, Eastwood 51, 1895 (CAS!). Dicoria wetherillii Eastw. Proc. Calif. Acad. II, 6: 299. 1896. Asteraceae. = D. brandegei Gray (monstrous form?) San Juan Co., San Juan River, Wetherill sn, 1895 (CAS!). Distichlis maritima var. laxa Holm Bot. Gaz. 16: 277. 1891. Poaceae. = D. stricta (Torr.) Rvdb. Utali, Lake Park, Tracy sn, 1887 (?). Dodecatheon zionense Eastw. Leafl. W. Bot. 2: 37. 1937. Primulaceae. = D. pulchcUitm (Raf.) Merr. Wash- ington Co., Zion Canyon, Eastwood & Howell 1144, 19.33 (CAS!). Draha apiculata C.L. Hitchc. Univ. Washington Publ. Biol. 11: 72. 1941. Bra.ssicaceae. = D. densifolia var. davisiae (C.L. Hitchc.) Welsh & Reveal Summit Co., Uinta Mts, Payson & Pavson 5048, 1926 (NY!;US!:RM!). Draha brachystylis Rydb. Bull. Torrey Bot. Club 29: 240. 1902. Brassicaceae. Salt Lake Co., Wasatch Mts., Alta, Jones 1.357, 1879 (NY!;US!;CAS!;RM!;POM!). Draha maguirei C.L. Hitchc. var. burkei C.L. Hitchc. Univ. Washington Publ. Biol. 11: 70. 1941 Brassicaceae. Box Elder Co., Cottonwood Canyon, Burke 2968, 19.32 (UTC!). Draba maguirei C.L. Hitchc. var. maguirei Univ. Washington Publ. Biol. 11: 70. 1941. Brassicaceae. Cache Co., Bear River Range, Maguire et al. 14161, 19.36 (WTU!;UTC!;NY!). Draba sobolifera Rydb. Bull. Torrey Bot. Club 30: 251. 1903. Brassicaceae. Syn: D. uncinalis Rydb. Piute Co., Tate Mine, Jones .59.36, 1894 (NY!;US!;POM!). Draba spectabilis Greene var. glabrescens Schulz Pflanzenr. 4. Fam. 105 (Heft 89) : 84. 1927. Brassicaceae. = D. spectabilis Greene San Juan Co., LaSal Mts., Walker 275, 1912 (NY!;US!). Draba subalpina Goodm. & Hitchc. Ann. Missouri Bot. Card. 19: 77. 19.32. Brassicaceae. Iron Co., Cedar Breaks, Goodman & Hitchcock 1622, 1930 (NY!;RM!). Draba uncinalis Rydb. Bull. Torrey Bot. Club 30: 251. 1903. Brassicaceae. = D. sobolifera Rydb. Piute Co., Tate Mine, Jones 5940am, 1894 (US!;POM!). Draba valida Goodding Bot. Gaz. 37: 55. 1904. Brassicaceae. = D. lanceolata Royle Uintah Co., Dyer Mine, Goodding 1402, 1902 (US!;RM!). Draba zionensis C.L. Hitchc. Univ. Washington Publ. Biol. 11: 49. 1941. Brassicaceae. = D. asprella var. zionensis (C.L. Hitchc.) Welsh & Reveal Washington Co., Zion Canyon, Jones sn, 1923 (POM!;CAS!;BRY!). Drymocallis micropetala Rydb. N. Amer. Fl. 22: 375. 1908. Rosaceae. = Potentilla glandulosa var. micropetala (Rydb.) Welsh & Johnst. Salt Lake Co., City Creek Can- yon, Rydberg 61,53, 1905 (US!;NY!). Echinocactus johnsonii Parry ex Engelm. Rep. U.S. Geol. Explor 40th Parallel, Bot. 5: 117. 1871. Cactaceae. = NeoUoydia johnsonii (Parrv) L. Benson Washington Co., near St. George, Johnson sn, 1870? (GH?). Echinocactus whipplei Engelm. & Bigel. var. spin- osior Engelm. Trans. Acad. Sci. St. Louis 2: 199. 1863. Cactaceae. = Sclerocactus pubispinus {Enge]m.) L. Ben- son Utah, desert valley west of Camp Floyd, H. Engel- mann sn, 18.59 (MO). Echinocereus engelmannii (Parry) Rumpler var. pur- pureus L. Benson Cact. & Succ. J. (Los Angeles) 41: 127. 1969. Cactaceae. Washington Co., near St. George, Ben- son 1.3637, 1949 (POM). Edwinia macrocalyx Small N. Amer. Fl. 22: 176. 1905. Saxifragaceae. = Jamesia americana T. & G. Utah Co., American Fork Canyon, Watson 371, 1869 (US!). Elymus salina Jones Proc. Calif. Acad. II, 5: 725. 1895. Poaceae. Sevier Co., Salina Pass, Jones 5447, 1894 (RM!;BRY!;US!;NY!). Emmenanthe foliosa Jones Zoe 4: 278. 1893. Hydro- phyllaceae. = PhaceUa salina (A. Nels.) J.T. Howell Tooele (?) Co.. Deep Creek, Jones sn, 1891 (POM!;US!). Encelia microcephala Gray Proc. Amer. Acad. 8: 657. 1873. Asteraceae. = Helianthella microcephala (Gray) Gray San Juan Co., Sierra .\bajo, Newberry sn, 1859 (NY!). Encelia nudicaulis Gray Proc. Amer. Acad. 8: 656. 1873. Asteraceae. = Enceliopsis nudicaulis (Gray) A. Nels. Utah, Bishop sn, 1872 (US!). Epilobium palmeri Rydb. Bull. Torrey Bot. Club 31: 569. 1904. Onagraceae. = E. saximontanum Hausskn. Washington Co., (s. Utah), Palmer 156, 1877 (NY!). Epilobium tracyi Rydb. Bull. Torrey Bot. Club 40: 63. 1913. Onagraceae. = E. hrachycarpiim Presl Weber Co., Ogden, Tracy & Evans 547, 1887 (NY!). Eremocarya muricata Rydb. Bull. Torrey Bot. Club 36: 677. 1909. Boraginaceae. = Cryptantha micrantha (Torr.) Johnst. Southern Utah, Parry 164, 1874 (NY!;1SC!). Erigeron abajoensis Cronq. Brittonia 6: 168. 1947. As- teraceae. San Juan Co., Abajo Mts., Rydberg & Garrett 97.55, 1911 (NY!;UT!). June 1982 Welsh: Utah Plant Types 167 Erigeron caespitosus Nutt. var. laccoliticus Jones Proc. Calif. Acad. II, 5: 696. 1895. Asteraceae. = E. caespitosus Nutt. Garfield Co., Henry Mts., Jones 5661, 1894 {POM!:US!). Erigeron caespitosus Nutt. var. laccoliticus Jones Proc. Calif. Acad. II, 5: 696. 1895. Asteraceae. = £. caespitosus Nutt. Garfield Co., Henry Mts., Jones 5661, 1894 (POM!;US!). Erigeron caespitosus Nutt. var. nauseosus Jones Proc. Calif. .\cad. II, 5: 696. 1895. Asteraceae. = E. nauseosus (Jones) Cronq. Piute Co., Marysvale, Jones 5386, 1894 (POM!;US!;NY!). Erigeron cinereus var. aridus Jones Proc. Calif. Acad. II, 5: 695. 1895. Asteraceae. = E. divergens T. & G. Washington Co., Silver Reef, Jones 5149v, 1894 (POM!;US!). Erigeron controversus Greene Leafl. Bot. Obs. & Grit. 2: 206. 1912. Asteraceae. = E. garrctti A. Nels. Salt Lake Co., Alta, Jones 1207, 1879 (US!;POM!). Erigeron cronquistii Maguire Brittonia 5: 201. 1944. Asteraceae. Cache Co., Logan Canyon, Maguire 16681, 1939 (NY!;US!;UTC!). Erigeron eatonii Gray Proc. Amer. Acad. 16: 91. 1880. Asteraceae. Summit Co., Uinta Mts., Watson 546, 1869 (US!:NY!). Erigeron eatonii Gray ssp. typicus Cronq. var. mo- lestus Cronq. Brittonia 6: 172. 1947. .Asteraceae. = £. eatonii Grav Tooele Co., Mt. Deseret Peak, Stansbury Range. Maguire and Holmgren 21773, 1943 (NY!;UTC!).' Erigeron flagellaris Gray var. trilobatus Maguire & Cronq. Brittonia 6; 258. 1947. Asteraceae. = £. prose- hjticus Nesom Iron Co., Cedar Breaks, Maguire 14947, 19.34 (NY!;UTC!). Erigeron fruticetorum Rydb. Fl. Rocky Mts. 906, 1067. 1917. Asteraceae. = E. formosissimus Greene San Juan Co., La Sal Mts., Rvdberg & Garrett 8912, 1911 (NY!). Erigeron garrettii A. Nels. in Coult. & Nels. Man. Bot. Centr. Rocky Mts. 526. 1909. Asteraceae. Syn: E. controversus Greene Salt Lake Co., Big Cottonwood Canyon, Garrett 1310, 1905 (RM!;UT!). Erigeron kachinensis Welsh & Moore Proc. Utah Acad. 45: 2.31. 1968. Asteraceae. San Juan Co, Natural Bridges, Welsh & Moore 2.398, 1963 (BRY!;NY!). Erigeron leiomerus Gray Syn. Fl. N. Amer. II. 1: 211. 1884. .\steraceae. Svn: E. minuscuhis Greene Summit Co., Uintas, Watson'504, 1869 (NY!). Erigeron leiophyllus Greene Leafl. Bot. Obs. & Grit. 2: 218. 1912. Asteraceae. = E. speciosus var. macr- anthus (Nutt.) Cronq. Salt Lake Co., Fort Douglas, Jones sn, 1880 (US!;POM!). Erigeron maguirei Cronq. Brittonia 6: 165. 1947. .As- teraceae. Emerv Co., Calf Spring Wa.sh, Maguire 18459, 1940 (NY!:UTC!). Erigeron mancus Rydb. Fl. Rocky Mts. 902, 1067. 1917. Asteraceae. Syn: £. pinnatisectus var. insoJens Macbr. & Pavson Grand Co., La Sal Mts., Rvdberg & Garrett 8671, 1911 (NY!;US!;UT!;RM:). Erigeron minusculus Greene Leafl. Bot. Obs. & Grit. 2: 8. 1909. .Asteraceae. = £. leiomcris Gray Salt Lake Co., Big Cottonwood Canyon, Garrett, 1906 (UT?). Erigeron pinnatisectus A. Nels. var. insolens Macbr. & Payson Contr. Gray Herb. II, 49: 79. 1917. Aste- raceae. = £. mancus Rvdb. Grand Co., La Sal Mts., Walker 271, 1912 (GH;MiNN;RM!). Erigeron pulvinatus Rydb. non Wedd. Fl. Rocky Mts. 911, 1067. 1917. Asteraceae. = £. compactus Blake Juab Co., Deep Creek, Jones sn, 1891 (US!;POM!;BRY!;NY!). Erigeron pumilus Nutt. ssp. concinnoides Cronq. var. subglaber Cronq. Brittonia 6: 183. 1947. Asteraceae. San Juan Co., Monticello, Rvdberg & Garrett 9141, 1911 (US!;RM!;UT!). Erigeron religiosus Cronq. Brittonia 6: 258. 1947. .As- teraceae. Washington Co., Clear Creek, Eastwood & Howell 6.3.39, 1938 (CAS!;US!). Erigeron sionis Cronq. Brittonia 6: 258. 1947. Aste- raceae. Washington Co., Zion National Park, Pilsbry .sn, 1925 (PHI). Erigeron sparsifolius Eastw. Proc. Calif. Acad. II, 6: 297. 1896. Asteraceae. = £. utahensis Gray var. spar- sifolius (Ea.stw.) Cronq. San Juan Co., Willow Creek, Eastwood 48, 1895 (CAS!). Erigeron stenophyllus D.C. Eaton Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot 5: 152. 1871. Asteraceae. = £. arenarioides (D.C. Eaton) Rydb. Salt Lake Co., Cotton- wood Canyon, Watson 547, 1869 (US!;NY!). Erigeron stenophyllus var. tetrapleuris Gray Proc. Amer. Acad. 8: 650. 1873. .Asteraceae. = £. utahensis Gray Kane Co., Kanab, Thompson sn, 1870 (GH). Erigeron uintahensis Cronq. Bull. Torrey Bot. Club 70: 270. 1943. Asteraceae. Summit Co., Mill Creek, Fay- son 4894. 19.36 (RM!;US!). Erigeron ursinus D.C. Eaton Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5; 148. 1871. Asteraceae. Summit Co., Bear River Canyon, Watson 5.34, 1869 (US!;NY!). Erigeron vagus Payson Univ. Wyoming Publ. Bot. 1: 179. 1926. Asteraceae. Grand Co., LaSal Mts., Payson & Payson 40.33, 1924 (US!;BRY!;RM!:NY!). Eriogonum ammophilum Reveal Phytologia 23: 163. 1972. Polvgonaceae. Millard Co., Ibex Warm Point, N. & P. Holmgren 46.50, 1970 (US!;NY!;BRY!;UTC!). Eriogonum aretioides Barneby Leafl. W. Bot. 5: 154. 1949. Polvgonaceae. Garfield Co., Widtsoe, Ripley & Barneby 8.570, 1947 (CAS!;NY!;BRY!). Eriogonum aureum Jones Proc. Calif. Acad. II, 5: 718. 1895. Polygonaceae. = £. corymbosum var. glutinosusm (Jones) Jones Washington Co., St. George, Jones 6091, 1894(POM!;US!;NY!).' Eriogonum batemanii Jones Contr. W. Bot. 11: 11. 1903. Polvgonaceae. Carbon Co., Price Valley, Jones sn, 1903(US!';POM!). Eriogonum bicolor Jones Zoe 4: 281. 1893. Polyg- onaceae. Grand Co., Thompsons Spring, Jones sn, 1891 (US!;POM!:NY!). Eriogonum biumbellatum Rydb. Bull. Torrey Bot. Club .39: .306. 1912. Polvgonaceae. = £. umbellatum var. subaridum Stokes Sevier Co., Fish Lake, Rydberg & Carlton 7409. 1905 (US!;RM!:NY!). Eriogonum brevicaule Nutt. var. pumilum Stokes in Jones Contr. W. Bot. 11: 10. 1903. Polygonaceae. = £. brevicaule var. laxifolium (T. & G.) Reveal Carbon Co., between Colton and Kyune, Jones 5603, 1894 (POM?). Eriogonum cernuum Nutt. var. tenue T. & G. Proc. Amer. Acad. 8: 182. 1870. Polygonaceae. = £. cernuum var. cernuum Weber Co., Weber Valley, Watson 10.36, 1869 (US!). Eriogonum cernuum Nutt. var. umbraticum Eastw. Proc. Calif. Acad. II, 6:: 319. 1896. Polygonaceae. = £. cen^uum var. cernuum San Juan Co., McElmo Creek, Eastwood sn, 1892 (CAS?). 168 Great Basin Naturalist Vol. 42, No. 2 Eriogonum chrysocephalum Gray ssp. alpestre Stokes Genus Eriogonum 93. 1936. Polygonaceae. = E. pan- guicense var. alpestre (Stokes) Reveal Iron Co., Cedar Breaks, Goodman & Hitchcock 1601, 1930 (CAS!; NY!;UTC!). Eriogonum clavellatum Small Bull. Torrey Bot. Club 25: 48. 1898. Polygonaceae. San Juan Co., Bartons range, Eastwood 1,32, 1895 (US!;NY!). Eriogonum confertiflorum var. stansburyi Benth. in DC Prodr. 14: 17. 1856. Polygonaceae. = E. brevicaiile var. brevicaule Utah (label data illegible), Stansbury sn, 1850 (NY!). Eriogonum corymbosum Benth. in DC. var. albogil- vum Reveal Great Basin Naturalist 27: 218. 1967. Polyg- onaceae. = E. X duchesnense Reveal Duchesne Co., In- dian Canyon, J & C Reveal 726, 1967 (US!;BRY!; NY!;UTC!). Eriogonum corymbosum Benth. in DC. var. davidsei Reveal Great Basin Naturalist 27: 216. 1967. Polyg- onaceae. Carbon Co., Wellington, Reveal & Davidse 956, 1967 (US!;BRY!;UT!;RM!;NY!;UTC!;ISC!). Eriogonum corymbosum Benth. in DC. var. divarica- tum T. & G. in Beckwith Rep. U.S. Explor. & Surv. R.R. Pacific 2: 29. 1857. Polygonaceae. Emery Co., Green River, Creutzfeldt sn, 1853 (NY!). Eriogonum corymbosum Benth. in DC. var. erectum Reveal & Brotherson in Reveal Great Basin Naturalist 27: 213. 1968. Polygonaceae. Wasatch Co., W Duch- esne, Holmgren & Reveal ,3022, 1966 (UTC!;US!; BRY!;UT!;NY!;ISC!). Eriogonum corymbosum Benth. in DC. var. mat- thewsiae Reveal Phytologia .35: 441. 1976. Polygonaceae. Washington Co., Springdale, Welsh et al. 9509, 1969 (BRY!;UT!). Eriogonum crispum L.O. Williams Bull. Torrey Bot. Club 59: 427. 19,32. Polygonaceae. = E. corymbosum var. ghitinosum (Jones) Jones Iron Co., Cedar Canyon, Garrett 6027, 1931 (RM!;UT!). Eriogonum cronquistii Reveal Madrono 19: 289. 1969. Polygonaceae. Garfield Co., Henry Mts., Holmgren & Reveal .3010, 1966 (BRY!;RM!;UT!;NY!;US!;UTC!;ISC!). Eriogonum deflexum Terr, in Ives ssp. hookeri var. gilvum Stokes Genus Eriogonum 45. 1936. Polygonaceae. = E. hookeri Wats. Davis Co., Antelope Island, Cottam 6407, 19,35 (UT!;UTC!). Eriogonum deflexum Torr. in Ives ssp. ultrum Stokes Genus Eriogonum 45. 1936. Polygonaceae. = £. nutans var. nutans Sevier Co., Sevier Valley, Eastwood & How- ell 623, 1933 (CAS!). Eriogonum duchesnense Reveal (hybrid) Great Basin Naturalist 27: 194. 1968. Polygonaceae. Syn: E. corymbo- sum var. albogilvum Reveal Duchesne Co., Indian Can- yon, Reveal 678, 1964 (UTC;US!;BRY!;NY!;UTC!). Eriogonum dudleyanum Stokes Genus Eriogonum 90. 1936. Polygonaceae. = £. kearneyi Tidestr. var. kearneyi Tooele Co., Skull Valley, Jones sn, 1896 (CAS!;UT!). Eriogonum effusum Nutt. var. durum Stokes Genus Eriogonum 80. 1936. Polygonaceae. = E. corymbosum var. corymbosum Carbon Co., Sunnyside, Jones 1113a, 1901 (POM!;NY!). Eriogonum effusum Nutt. ssp. orbiculatum Stokes Genus Eriogonum 79. 1936. Polygonaceae. = E. co- rymbosum var. orbiculatum (Stokes) Reveal & Broth- erson Emery Co., Green River, Jones sn, 1915 (POM!;BRY!). Eriogonum effusum Nutt. ssp. pallidum var. shandsii Stokes Genus Eriogonum 81. 1936. Polygonaceae. = E. leptocladon var. leptocladon San Juan Co., 3 mi s Indian Creek, Shands 1,30, 1931 (UT!). Eriogonum ephedroides Reveal Madrono 19: 295. 1969. Polygonaceae. Uintah Co., 10 mi s Bonanza, Hol- mgren et al. 2265, 1965 (UTC!;BRY!;UT!;RM!;ISC!). Eriogonum eremicum Reveal Phytologia 23: 165. 1972. Polygonaceae. Millard Co., se Garrison, Holmgren et al. 2247, 1965 (UTC!;BRY!;UT!;RM!;ISC!). Eriogonum filicaule Stokes Genus Eriogonum 32. 1936. Polygonaceae. = £. subreniforme Reveal Wash- ington Co., Springdale, Eastwood & Howell 1171, 1933 (CAS!;US!;NY!). Eriogonum filiforme L.O. Williams Bull. Torrey Bot. Club 59: 428. 1932. Polygonaceae. = £. wetherillii Eastw. Wayne Co., w Hanksville, Garrett 5975, 1931 (UT!;RM!). Eriogonum flexum Jones var. ferronis Jones Contr. W. Bot. 11: 15. 1903. Polygonaceae. = £. flexum Jones Emery Co., w Perron, Jones 5454, 1894 (US!;POM!). Eriogonum friscanum Jones Contr. W. Bot. 11: 14. 1903. Polygonaceae. = £. microthecum var. foliosum (T. & G.) Reveal Beaver Co., Frisco, Jones (POM?). Eriogonum grayii Reveal Phytologia 25: 193. 1973. Polygonaceae. Salt Lake Co., Lake Blanche, Holmgren et al. 7121, 1947 (UTC!;BRY!;ISC!). Eriogonum heracleoides Nutt. var. utahense Gandg. Bull. Soc. Bot. Belgium 42: 190. 1906. Polygonaceae. = £. heracleoides Nutt. var. heracleoides Cache Co., Lin- ford sn, 1897 (US!). Eriogonum hookeri Wats. Proc. Amer. Acad. 14: 295. 1879. Polygonaceae. Syn: £. deflexum ssp. hookeri var. gilvum Stokes Utah Co., American Fork Canyon, Wat- son 10,33, 1869 (US!;NY!). Eriogonum howellianum Reveal Phytologia 25: 204. 1973. Polygonaceae. Syn: £. glandidosum authors, not (Nutt.) Nutt. Millard Co, se Garrison, Holmgren et al 2248, 1965 (US!;BRY!;UT!;RM!;UTC!). Eriogonum humivagans Reveal Madrono 19: 219. 1969. Polygonaceae. San Juan Co., 13.5 miles e Mon- ticello, Holmgren & Reveal 3001, 1969 (UTCi; US!;RM!;BRY!;UT!;NY!;ISC!). Eriogonum hylophilum Reveal & Brotherson Great Basin Naturalist 27: 190. 1968. Polygonaceae. Duchesne Co, Gate Canyon, Holmgren & Reveal 3017, 1966 (UTC!;US!;RM!;BRY!;UT!;NY!;ISC!). Eriogonum insigne Wats. Proc. Amer. Acad. 14: 295. 1879. Polygonaceae. Iron Co., Red Creek, Palmer 431, 1877 (NY!;BRY!;ISC!). Eriogonum intermontanum Reveal Madroiio 19: 293. 1969. Polygonaceae. Grand Co., Middle Canyon, Roan Cliffs, Holmgren et al. 2278, 1965 (UTC!;BRY!; UT!;RM!;ISC!). Eriogonum jamesii Benth in DC. var. rupicola Reveal Phytologia 25: 202. 1973. Polygonaceae. Washington Co., Zion National Park, Reveal & Reveal 2874, 1972 (US!;BRY!;NY!;UTC!). June 1982 Welsh: Utah Plant Types 169 Eriogonum kearneyi Tidestr. Proc. Biol. Soc. Wash- ington 26: 122. 1913. Polygonaceae. Syn: E. dtidleyanum Stokes Tooele Co., w Tooele, Kearney & Shantz 3218 (US!). Eriogonum kingii var. laxifolitim T. & G. Proc. Anier. Acad. 8: 165. 1870. Polygonaceae. = £. brevicaule Nutt. var. kixifoUum (T. & G.) Reveal Salt Lake Co., Parleys Peak, Watson 1021, 1869 (US!). Eriogonum lancifolium Reveal & Brotherson in Re- veal Great Basin Naturalist 27: 188. 1967. Polygonaceae. Carbon Co., e Wellington, Reveal & Davidse 955, 1967 (UTC!;US!;RM!;BRY!;UT!;NY!;ISC!). Eriogonum leptocladon T. & G. in Beckwith var. lep- tocladon Rep. U.S. Explor. & Surv. R.R. Pacific 2: 129. 1857. Polygonaceae. Syn: E. effusum spp. pcdlidum var. shandsii Stokes Einerv Co., Green River, Creuzfeldt sn, 1853 (NY!). Eriogonum hganum A. Nels. Bot. Gaz. 54: 149. 1912. Polygonaceae. Cache Co., Logan, Smith 1704, 1909 (RM!;BRY!). Eriogonum longilobum Jones Proc. Calif. Acad. II, 5: 720. 1895. Polygonaceae. = E. shockleyi var. longilobum (Jones) Reveal Carbon Co., near Price, Jones 5590, 1894 (POM!;US!). Eriogonum medium Rydb. Fl. Rocky Mts. 220, 1061. 1917. Polvgonaceae. = E. brevicaule Nutt. var. laxifo- lium (T. & G.) Reveal Juab or Utah Co., Mt. Nebo, Ryd- berg sn, 1905 (US!;RM!;NY!). Eriogonum nanum Reveal Phytologia 25: 194. 1973. Polygonaceae. Box Elder Co., Willard Peak, Reveal & Holmgren 665, 1964 (UT!;BRY!;RM!). Eriogonum natum Reveal in Welsh, Atwood, & Re- veal Great Basin Naturalist 35: 363. 1975. Polygonaceae. Millard Co, 43 mi w Delta, Reveal & Reveal 3924, 1975 (BRY!;NY!;UTC!). Eriogonum nehonii L.O. Williams Bull. Torrey Bot. Club 59: 428. 19'37. Polygonaceae. = £. microtheciim Nutt. var. foliosum (Torr.) Reveal San Juan Co., Geyser Basin, Walker .368, 1912 (RM!;UT!). Eriogonum nudicaule Small ssp. garrettii Stokes Genus Eriogonum 8.3. 19.36. Polygonaceae. = E. brevi- caule Nutt. var. brevicaule Summit Co., Echo Reservoir, Garrett 7068, 19.35 (US!;UT!;UTC!). Eriogonum nudicaule Small ssp. ochroflorum Stokes Genus Eriogonum 83. 1936. Polygonaceae. = E. spa- thulatum Gray Sevier Co., Clear Creek Canyon, Jones 6104, 1894 (p6m!;BRY!). Eriogonujn nudicaule Small ssp. parleyense Stokes Genus Eriogonum 84. 19.36. Polygonaceae. = E. brevi- caule Nutt. var. brevicaule Salt Lake Co., Parleys Can- yon, Stokes 213, 1934 (CAS!;NY!;UTC!;BRY!). Eriogonum nummulare Jones Contr. W. Bot. 11: 13. 1903. Polvgonaceae. Tooele Co., Dutch Mt., Jones sn (POM?). Eriogonum ochrocephalum var. angustum Jones Contr. W. Bot. 11: 9. 1903. Polygonaceae. = £. brevi- caule Nutt. var. laxifolium (T. & G.) Reveal Tooele Co., Johnsons Pass, Jones sn (POM?). Eriogonum ostlundii Jones Contr. W. Bot. 11: 12. 1903. Polvgonaceae. Sevier Co., near Elsinore, Jones sn (POM?). ' Eriogonum ovalifolium Nutt. var. utahense Gandg. Bull. Soc. Bot. Belgium 42: 191. 1906. Polygonaceae. = £. ovalifolium Nutt. var. ovalifolium Cache Co., Linford sn, 1897 (US!). Eriogonum palmeri Wats. Proc. Amer. Acad. 14: 267. 1879. Polygonaceae. = £. plumatella Dur. & Hilg. Washington (?) Co., Palmer sn, 1870 (US!). Eriogonum parryi Gray Proc. Amer. Acad. 10: 77. 1874. Polygonaceae. = £. brachypodum T. & G. Wash- ington Co., St. George, Parry 239, 1874 (US!;NY!;BRY!). Eriogonum pauciflorum var. panguicense Jones Contr. W. Bot. 11: 9. 1903. Polygonaceae. = £. pan- guicense (Jones) Reveal Garfield Co., Panguitch, Jones sn, 1890 (POM!). Eriogonum pharnaceoides Torr. in Sitgr. var. cervi- num Reveal Great Basin Naturalist 34: 245. 1974. Polyg- onaceae. Washington Co., Pine Valley Mts., Atwood & Higgins 5895, 1973 (US!;BRY!;RM!;UTC!). Eriogonum porteri Small Bull. Torrey Bot. Club 25: 41. 1898. Polygonaceae. = £. umbellatum Torr. var. porteri (Small) Stokes Summit Co., Bear River Canyon, Watson 1014, 1869 (US!;NY!). Eriogonum puberulum Wats. Proc. Amer. Acad. 14: 295. 1879. Polygonaceae. Iron Co., Red Creek, Palmer 429, 1877 (US!;NY!;BRY!;ISC!). Eriogonum pulvinatum Small Bull. Torrey Bot. Club 25: 44. 1898. Polygonaceae. = £. shockleyi Wats. var. shockleyi Beaver Co., Milford, Jones 1775, 1880 (US!;UT!;POM!;NY!;UTC!). Eriogonum ramosissimum Eastw. Proc. Calif. Acad. II, 6: 322. 1896. Polygonaceae. = £. leptocladon T. & G. var. ramosissiinum (Eastw.) Reveal San Juan Co., Butter Spring, Eastwood 134, 1895 (US!;CAS!). Eriogonum revealianum Welsh Great Basin Naturalist 30: 17. 1970. Polygonaceae. = £. corymbosum var. re- vealianum (Welsh) Reveal Garfield Co., s Antimony, Welsh & Welsh 9389, 1969 (BRY!;US!;UT!;RM!;NY!). Eriogonum rubiflorum Jones Zoe 4: 281. 1893. Polyg- onaceae. = E. nutans T. & G. var. nutans Tooele Co., Dugway, Jones sn, 1891 (POM!;US!;NY!). Eriogonum saurinum Reveal Great Basin Naturalist 27: 197. 1967. Polygonaceae. Uintah Co., 10 mi e Vernal, Holmgren & Reveal 3019, 1966 (UTC!;UT!;BRY!;US!;RM!;NY!;ISC!). Eriogonum scabrellum Reveal Ann. Missouri Bot. Card. 55: 74. 1968. Polygonaceae. Grand Co, West- water, Reveal & Davidse 949, 1967 (UTC!;US!;BRY!;UT!;RM!;NY!;ISC!). Eriogonum smithii Reveal Great Basin Naturalist 24: 202. 1967. Polygonaceae. Emery Co., San Rafael Desert, Holmgren &' Reveal .30i2, 1966 (UTC!:US!; BRY!;UT!;RM!;ISC!). Eriogonum soredium Reveal Great Basin Nat. 41: 229. 1981. Polygonaceae. Beaver Co., Grampian Hill, near Frisco, Welsh et al. 20192,1980 (BRY!). Eriogonum spathuliforme Rydb. Bull. Torrey Bot. Club 39: 307. 1912. Polygonaceae. = E. ostlundii Jones Piute Co., near Mt.Belknap, Stokes sn, 1900 (US!). Eriogonum spathulatum Gray Proc. Amer. .\cad. 10: 76. 1874. Polygonaceae. Syn: £. nudicaule Small ssp. ochroflorum Stokes Sevier (?) Co., Sevier River Valley, Parry 245, 1874 (ISC!). Eriogonum subreniforrne Wats. Proc. Amer. Acad. 12: 260. 1877. Polygonaceae. Syn: £. filicaide Stokes Wash- ington Co., Parry 237, 1874 (ISC!). Eriogonum sulcatum Wats. Proc. Amer. Acad. 14: 296. 1879. Polygonaceae. = £. heennanii var. sulcatum (Wats.) Munz & Reveal Washington Co., near St. George, Palmer 432, 1877 (US!;NY!;BRY!). 170 Great Basin Naturalist Vol. 42, No. 2 Eriogonum sulcatum Wats. var. argense Jones Contr. W. Bot. 11: 15. 1903. Polygonaceae. = E. heermannii Dur. & Hilg. var. argense (Jones) Munz Utah, Jones. Eriogonum tenellum ssp. cottamii Stokes Genus Eriogonum 70. 1936. Polygonaceae. = E. brevicaiile var. cottamii (Stokes) Reveal Utah Co., West Mountain, Cot- tarn 411, 1925(BRY!;UT!). Eriogonum tenellum var. grandiflorum Gandg. Bull. Soc. Bot. Belgium 42: 197. 1906. Polygonaceae. = E. mi- crothecum var. laxiflorum Hook. Rich Co., Linford sn, 1897 (?). Eriogonum thompsonae Wats. var. albiflorum Reveal Madroiio 19: 299. 1969. Polygonaceae. Washington Co., w Virgin, Holmgren & Reveal 2991, 1966 (UTC!; US!;BRY!;UT!;RM!;NY!;ISC!). Eriogonum thompsonae Wats. var. thompsonae Amer. Naturalist 7: .302. 1873. Polygonaceae. Kane Co., Kanab, Thompson sn, 1872 (GH;US!;BRY!;NY!). Eriogonum triste Wats. Proc. Amer. Acad. 10: 347. 1875. Polygonaceae. = E. alatum Torr. in Sitgr. Kane Co., Siler sn, 1873 (US!;NY!;BRY!;ISC!). Eriogonum umbellatum Torr. var. desereticum Reveal in Welsh, Atwood, & Reveal Great Basin Naturalist 35: .365. 1975. Polygonaceae. Utah Co., Mt. Timpanogos, near Timpooneke Campground, Reveal 3702. 1974 (US!;BRY!;NY!;UTC!). Eriogonum umbellatum Torr. var. glabratum Stokes Genus Eriogonum 109. 1936. Polygonaceae. = E. um- bellatum Torr. var. aureum ( Gandg.) Reveal Emery Co., Huntington Canyon, Garrett 7036, 1935 (UT!). Eriogonum villiflorum Gray var. tumulosum Barneby Leafl. W. Bot. 5: 153. 1949. Polygonaceae. = £. tumulo- sum (Barneby) Reveal Emery Co., sw Woodside, Ripley & Barneby 8678, 1947 (CAS!;NY!;BRY!;UTC!). Eriogonum viridulum Reveal Proc. Utah Acad. 42: 287. 1966. Polygonaceae. Duchesne Co., 8 mi e Duch- esne, Reveal 675, 1964 (UTC!;BRY!;UT!;RM!;NY!;ISC!). Eriogonum wasatchense Jones Contr. W. Bot. 11: 11. 1903. Polygonaceae. = £. brevicaule var. wasatchense (Jones) Reveal Utah Co., American Fork Canyon, Jones 1877, 1880(US!;POM!;UTC!). Eriogonum wetherillii Eastw. Proc. Calif. Acad. II, 6: 319. 1896. Polygonaceae. Syn: E. filiforme L.O. Williams San Juan Co., San Juan River, Eastwood 124, 1895 (CAS!). Eriogonum zionis J. T. Howell Leafl. W. Bot. 2: 253. 1940. Polygonaceae. Washington Co, Zion National Park, Eastwood & Howell 6344, 1938 (CAS!). Eritrichium barbigerum Gray Synop. Fl. N. Amer. 2(1): 194. 1878. Boraginaceae. = Cryptantha barbigera (Gray) Greene Washington Co., St. George, Parry 171, 1874 (GH;ISC!). Eritrichium elongatum Wight var. paysonii Johnst. J. Arnold Arb. 33: 67. 1952. Boraginaceae. = £. nanum var. elongatum (Wight) Cronq. Summit Co., Uinta Mts, Maguire et al 14385, 19.36 (RM!). Eritrichium holopterum Gray Proc. Amer. Acad. 12: 81. 1876. Boraginaceae. = ? Southern Utah, Bishop sn, 1873 (GH?). Eritrichium holopterum Gray var. submolle Gray Proc. Amer. Acad. 13: 374. 1878. Boraginaceae. = Cryp- tantha utahensis (Gray) Greene Washington Co., St. George, Palmer sn, 1870 (NY!). Eritrichium pterocaryum Torr. var. pectinatum Gray Proc. Amer. Acad. 10: 61. 1874. Boraginaceae. = Cryp- tantha pterocarya var. pterocarya Southern Utali, Parry sn, 1874 (NY!). Eritrichium setosissimum Gray Proc. Amer. Acad. 12: 80. 1876. Boraginaceae. = Cryptantha setosissima (Gray) Payson Sevier Co., Fish Lake, Ward sn, 1875 (GH). Erysimum asperum (Nutt.) DC. var. purshii Durand Trans. Amer. Phil. Soc. II. 159. 1860. Brassicaceae. Syn: E. capitatum (Dougl.) Greene Salt Lake Co., near Great Salt Lake, Salt Lake Valley, Carrington sn, 1857 (P). Erythrocoma brevifolia Greene Leafl. Bot. Obs. & Grit. 1: 176. 1906. Rosaceae. = Geum triflorum Pursh Garfield Co., Panguitch Lake, Jones 6002q, 1894 (US!). Erythronium utahense Rydb. Fl. Rocky Mts. 165, 1061. 1917. Liliaceae. = E. grandiflorum Pursh Salt Lake Co., Salt Lake, Stansbury sn, 1850 (NY!). Eschscholzia ludens Greene Pittonia 5: 272. 1905. Pa- pa veraceae. = E. minutiflora Wats. Washington Co., St. George, Jones 5110a, 1894 (US!;POM!). Euphorbia nephradenia Barneby Leafl. W. Bot. 10: 314. 1966. Euphorbiaceae. Kane Co., Cottonwood Can- yon, Barneby 14434, 1966 (NY!;US!;BRY!;UTC!). Euphorbia parryi Engelm. in Parry Amer. Naturalist 9: .350. 1875. Euphorbiaceae. Wa.shington Co., near St. George, Parry s47, 1874 (MO). Euphorbia podagrica I.M. Johnston Univ. Calif. Publ. Bot. 7: 440. 1922. Euphorbiaceae. Utah, Purpus 6432, 1898 (US!). Euphorbia robusta (Engelm.) Small ex Britt. & Brown var. interioris J.B.S. Norton N. Amer. Euphorbia 49. 1899. Euphorbiaceae. = E. robusta (Engelm.) Small Wasatch Mts., Watson 1081, 1869 (US!). Eurotia subspinosa Rydb. Bull. Torrey Bot. Club 39. 313. 1912. Chenopodiaceae. = Ceratoides subspinosa (Rydb.) J.T. Howell Washington Co., St. George, Goodd- ing 810, 1903 (US!). Festuca brevifolia R. Br. var. utahensis St-Yves Can- dollea 2: 257. 1925. Poaceae. = F. ovina L. Salt Lake Co., Alta, M.E.D. (possibly M.E.Jones) sn, 1895 (NY!). Festuca dasyclada Hack, ex Beal Grasses N. Amer. 2: 602. 1896. Poaceae. Emery (?) Co., Parry 93, 1874 (US!;ISC!;NY!). Festuca jonesii Vasey Contr. U.S. Natl. Herb. 1: 278. 1893. Poaceae. = F. subukita Trin. Salt Lake Co., City Creek Canyon, Jones sn, 1880 (US!;POM!). Fragaria virginiana Duchesne var. glauca Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 280. 1871. Ro- saceae. = F. virginiana Duchesne Summit Co., Parleys Park, Watson .322, 1869 (US!;NY!). Frasera albomarginata Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 220. 1871. Gentianaceae. = Sivertia albomarginata (Wats.) Kuntze Washington Co., near St. George, Palmer, 1879 (US!;NY!). Frasera utahensis Jones Zoe 2: 14. 1891. Gen- tianaceae. = Swertia paniculata (Torr.) St. John Kane (?) Co., (possibly House Rock Valley), Buckskin Mts., Jones sn, 1890 (US!;POM!). Fraxintts anomala Torr. in Wats. Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot. 5: 283. 1871. Oleaceae. San Juan (?) Co., Labrynth Canyon, Newberry sn, 1859 (NY!). Fritillaria dichroa Gandg. Bull. Soc. Bot. France 66: 291. 1920. Liliaceae. = F. pudica (Pursh) Spreng. Cache Co., Linford sn, 1897 (?). June 1982 Welsh: Utah Plant Types 171 Fritillaria leucella Gaiidg. Bull. Soc. Bot. France 66: 291. 1920. Liliaceae. = F. pudicci (Pursh) Spreng. Cache Co., Linford sii. 1897 (?). Fritillaria utahensis Gandg. Bull. Soc. Bot. France 66: 291. 1920. Liliaceae. = F. pticUca (Pursh) Spreng. Cache Co., Linford sn, 1897 (?). Gaillardia acaulis Gray Proc. Anier. Acad. 10: 73. 1874. Asteraceae. = G. parnji Greene Washington Co., St. George, Parrv 120, 1874 (NY!;NDG!). Gaillardia crassifolia Nels. & Macbr. Bot. Gaz. 61: 46. 1916. Asteraceae. = G. pinnatifida Torr.? Washing- ton Co., LaVerkin, Jones 5177, 1894 (US!;NY!;BRY!). Gaillardia flava Rydb. N. Amer. Fl. .34: 139. 1915. -Asteraceae. Emerv Co., Lower Crossing, Jones 6412, 1898(US!;BRY!;POM!). Gaillardia gracilis A. Nels. Bot. Gaz. 37: 276. 1904. Asteraceae. = G. pinnatifida Torr. Washington Co., Di- amond Valley, Goodding 894, 1902 (US!;RM!). Gaillardia spathulata Gray Proc. Amer. Acad. 12: 59. 1876. Asteraceae. Wavne Co.', Rabbit Valley, Ward 401, 1875(US!;NY!). Gaillardia straminea A. Nels. Univ. Wyoming Publ. Bot. 1: 137. 1926. Asteraceae. = G. pinnatifida Torr. Washington Co., LaVerkin, Jones 5177, 1894 (RM!;\Y!:BRY!). Galium bifolium Wats. Rep. U.S. Geol. E.\plor. 40th Parallel, Bot. 5: 134. 1871. Rubiaceae. Salt Lake Co. (?), Wasatch Mts., Watson 480, 1869 (US!;NY!). Galium desereticum Dempst. & Ehrend. Brittonia 17: 314. 1965. Rubiaceae. = G. mtiltiflonnn var. multi- florum Juab Co., Fish Springs, Jones sn, 1891 (US!;POM!). Galium filipes Rydb. Fl. Rocky Mts. 809, 1066. 1917. Rubiaceae. = G. mexicanum var. asperuhim Gray Salt Lake Co., Citv Creek Canyon, Leonard 16665, 1883 (NY!). Galium hypotrichium Gray ssp. scabriusculum Eh- rend. Contr. Dudley Herb. 5: 13. 1956. Rubiaceae. = G. multiflorum var. multiflorum Emery Co., Calf Spring Wash, Maguire 18437, 1940 (CAS!;Utc!). Galium hypotrichium ssp. utahense Ehrend. Contr. Dudley Herb. 5: 12. 1956. Rubiaceae. = G. multiflorum var. multiflorum Utah Co., Provo, Goodding 1116, 1902 (ISC!). Galium multiflorum Kellogg var. watsonii Gray Syn. Fl. N. Amer. 1(2): 40. 1884. Rubiaceae. = G. multi- florum var. multiflorum Syn: G. ivatsonii (Grav) Heller Wasatch Mts., Watson 484, 1869 (US!;NY!). Galium scabriusculum (Ehrend.) Dempst. & Ehrend. ssp. protoscabriusculum Dempst. & Ehrend. Brittonia 17: 312. 1965. Rubiaceae. = G. multiflorum var. multi- florum Carbon Co.. Castle Gate, Ehrendorfer & Stutz 5954. 1959(RM!:UTC!). Galium utahense Eastw. Leafl. W. Bot. 1: 55. 1933. Rubiaceae. = G. horeale L. Wasatch Co., Soldier Sum- mit, Eastwood 7668. 1918 (CAS!). Gentiana calycosa Griseb. ssp. apetala Maguire Madrono 6: 151. 1942. Gentianaceae. = G. calycosa Griseb. Duchesne Co., w Mt. .\gassiz, Maguire et al. 4225. 19.33 (UTC!). Gentiana tortuosa Jones Proc. Calif. .\cad. II, 5: 707. 1895. Gentianaceae. = Gentianella tortuosa (Jones) Gil- lett Garfield Co., Panguitch Lake, Jones 6008, 1894 (POM!:US!). Geranium marginale Rydb. ex Hanks & Small N. Amer. Fl. 25: 16. 1907. Geraniaceae. Garfield Co., Aquarius Plateau, Rydberg & Carlton 7401, 1905 (NY!). Gilia aggregata (Pursh) Spreng. ssp. aggregata var. attenuata f. utahensis Brand Pflanzenreich 4. Fani. 250: 116. 1907. Polemoniaceae. = G. aggregata var. riiacrosi- phon Kearney & Peebles Salt Lake Co., Alta, Jones 1122, 1879 (POM !;UTC!). Gilia arenaria Benth. var. rubella Brand in Engler Pflanzenreich 4. Fam. 250: 103. 1907. Polemoniaceae. = G. hutchinsifolia Rydb. Washington Co., St. George, Jones 1651, 1880 (Us'!;POM!;NY!). Gilia caespitosa Gray Proc. Amer. Acad 12; 80. 1876. Polemoniaceae. Wayne Co., Rabbit Valley, Ward 575, 1875 (US!;NY!;BRY!). Gilia congesta Hook. var. nuda Eastw. Proc. Calif. Acad. II, 6: 308. 1896. Polemoniaceae. = G. roseata Rvdb. Syn: G. nuda (Eastw.) Rydb. San Juan Co., Wil- low Creek, Eastwood 80, 1895 (US!;CAS!). Gilia congesta Hook. var. paniculata Jones Proc. Ca- lif. Acad. II, 5: 712. 1895. Polemoniaceae. = G. congesta var. congesta Emery Co., Huntington, Jones 5464m, 1894 (POM!;US!). Gilia debilis Wats. Amer. Naturalist 7: .302. 1873. Po- lemoniaceae. = Collomia debilis (Wats.) Greene Salt Lake Co., Wasatch Mts., above Salt Lake City, Wheeler sn, 1872 (US!). Gilia depressa Jones in Gray Proc. Amer. Acad. 16: 106. 1880. Polemoniaceae. Syn: Ipomopsis depressa (Jones) V. Grant Millard Co., Deseret, Jones 1772, 1880 (GH;US!:CAS!;POM!;BRY!;NY!;UTC!). Gilia filiformis Parry ex Gray Proc. Amer. Acad. 10: 75. 1874. Polemoniaceae. Washington Co., near St. George, Parry 187, 1874 (NY!;ISC!;BRY!). Gilia floribunda var. arida Jones Proc. Calif. Acad. II, 5: 713. 1895. Polemoniaceae. = Leptodactylon wat- sonii (Gray) Rydb. Wayne Co., Capitol Wash, Jones 5701a, 1894(POM!;US!).' Gilia frutescens Rydb. Bull. Torrey Bot. Club 40: 471. 1913. Polemoniaceae. = G. congesta var. frutescens (Rvdb.) Cronq. Washington Co., Springdale, Jones 5247, 1894 (NY!;US!). Gilia gracilis Hook. ssp. spirillifera var. nana Brand in Engler Pflanzenr. 4. Fam. 250: 92. 1907. Polemo- niaceae. = Microsteris gracilis (Hook.) Greene Millard Co., Fillmore, Jones 1684 (POM!;UT!). Gilia inconspicua (J.E. Sm.) Sweet ssp. eu- incotispicua var. variegata Brand in Engler Pflanzenr. 4. Fam. 250: 105. 1907. Polemoniaceae. = G. inconspic- ua (J.E. Sm.) Sweet Kane Co., Kanab, Jones 5280, 1894 Gilia latifolia Wats, in Parry Amer. Naturalist 9: 347. 1875. Polemoniaceae. Washington Co., near St. George, Parry 188, 1874 (GH;US!;CAS!;NY!;BRY!;ISC!). Gilia leptomeria Gray Proc. Amer. .-Kcad. 8: 278. 1870. Polemoniaceae. Syn: G. leptomeria var. tridentata Jones; G. triodon Eastw. Tooele Co., Stansbury Island, Watson 27, 1869 (NY!). Gilia leptomeria Gray var. tridentata Jones Proc. Ca- lif. Acad. II, 5: 713. 1895. Polemoniaceae. = G. lepto- meria Gray Emery Co., Emery, Jones 5445n, 1894 (POM;US!).' 172 Great Basin Naturalist Vol. 42, No. 2 Gilia incvickerae Jones Proc. Calif. Acad. II, 5: 712. 1895. Polemoniaceae. = G. pinnatifida Nutt. Piute Co., Marysvale, Jones 5-378, 1894 (POi\1!;US!;CAS!; BRY!:RM!;NY!). Gilia scoptilontm Jones Bull. Torrey Bot. Club 8; 70. 1881. Polemoniaceae. Syn: G. scopulorum var. deformis Brand Washington Co., St. George, Jones 1659, 1880 (US!;POM!;NY!;UTC!). Gilia scopulorum Jones var. deformis Brand Pflan- zenr. 4. Fani. 2.50; 109. 1907. Polemoniaceae. = G. sco- pulonnn Jones? .Southern Utiili, Parry 198, 1874 (?). Gilia stenothrysa Gray Proc. Amer. Acad. 8: 276. 1870. Polemoniaceae. Duchesne (?) Co., "Among cedars between Duchesne and Lake Fork," Fremont sn, 1845 (NY!). Gilia straminea Rydb. Bull. Torrev Bot. Club 40: 472. 1913. Polemoniaceae. = G. inconspicita var. sinuata (Hook.) Gray Washington Co., St. George, Palmer 325, 1877 (NY!). ' Gilia superba Eastw. Zoe 4: 123. 1893. Polemo- niaceae. = G. siihnuda Torr. San Juan Co., Hatches Wash, Eastwood sn, 1892 (CAS!;POM!). Gilia tenerrima Gray Proc. Amer. Acad. 8: 277. 1870. Polemoniaceae. Summit Co., Bear River Vallev, Wat.son 922, 1869 (NY!). Gilia tenuituba Rydb. Bull. Torrey Bot. Club 40: 472. 1913. Polemoniaceae. = G. aogregata var. macwsiphon Kearney & Peebles Beaver (?) Co., Beaver City, Palmer 320, 1877 (NY!;US!). Gilia tridactyla Rydb. Fl. Rocky Mts. 692,1065. 1917. Polemoniaceae. Piute Co., Brigham Peak, Jones .5949, 1894(US!;NY!;POM!). Gilia triodon Eastw. Zoe 4: 121. 1893. Polemo- niaceae. = G. leptomeria Gray San Juan Co., Ruin Can- yon. Eastwood sn, 1892 (CAS!). Gilia watsonii Gray Proc. Amer. Acad. 8: 267. 1870. Polemoniaceae. = Leptodactyhn watsonii (Gray) Rydb. Salt Lake Co., Cottonwood Canyon, Watson sn, 1869 (NY!). Glycosma maxima Rydb. Bull. Torrev Bot. Club 40: 67. 1913. Apiaceae. = Osmorhiza occidentalis (Nutt.) Torr. Juab Co., Mt. Nebo, Rydberg & Carlton 7585, 1905 (NY!). Glyptopleura setulosa Gray Proc. Amer. .Acad. 9: 211. 1874. Asteraceae. Washington Co., St. George, Palmer 6, 1870 (US!;NY!;BRY!). Grayia brandegei Gray Proc. Amer. Acad. 11: 101. 1876. Chenopodiaceae. San Juan (?) Co., San Juan R., Brandegee sn, 1875 (GH), Grindelia laciniata Rydb. Fl. Rocky Mts. 848, 1066. 1917. Asteraceae. = G. fastigiata Greene San Juan Co., Montezuma Canyon, Rydberg & Garrett 9692, 1911(NY!;USI). Grindelia stylosa Eastw. Proc. Calif. Acad. II, 6: 293. 1896. Asteraceae. = Vanclevea stylosa (Eastw.) Greene San Juan Co., Bartons range, Eastwood 713, 1895 (US!;CAS!;NY!). Gutierrezia sarothrae (Pursh) Britt. & Rusby var. pomariensis Welsh Great Basin Naturalist .30: 19. 1970. Asteraceae. = Xantliocephahim sarothrae var. poma- riensc (Welsh) Welsh Uintah Co., Orchard Creek Draw, Welsh etal. 9471, 1969 (BRY!). Cymnolomia his])ida Robins. & Greenm. var. ciliata Robins. & Greenm. Proc. Boston Soc. Nat. Hist. 29: 93. 1899. .Asteraceae. = Heliomeris hispida (Grav) Cockerell Syn: Viguiera ciliata (Robins. & Greenm.) Blake South- ern Utah, Palmer 245, 1877 (US!). Gymnolomia linearis Rydb. Bull. Torrey Bot. Club 37: 327. 1910. Asteraceae. = Heliomeris multiflora var. nevadensis (A. Nels.) Yates Washington Co., St. George, Palmer 241, 1877 (NY). Gymnolomia multiflora var. annua Jones Proc. Calif. ,\cad. II, 5: 698. 1895. Asteraceae. = Heliomeris long- ifolia var. annua (Jones) Yates Utah?, Jones ?, (?). Hackelia ibapensis Shultz & Shultz Brittonia 33: 157. 1981. Boraginaceae. Juab Co., Deep Creek Range, L. & J. Shultz 4:^50, 1980 (NY!;UTC!). Hackelia patens (Nutt.) Johnst. var. harrisonii Gentry Southw. Naturalist 19: 140. 1974. Boraginaceae. Wash- ington Co., Pine Valley, Gentry 2002, 1968 (NY!; CAS!;BRY!:UTC!). Halostachys occidentalis\ Wats. Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot. 5: 293. 1871. Chenopodiaceae. = AUcnrolfia occidentalis (Wats.) Kuntze Box Elder Co., Raft River Valley, Watson 995, 1869 (US!). Hamosa atratiformis Rydb. Bull. Torrey Bot. Club 34: 48. 1907. Fabaceae. = Astragalus straturensis Jones Wahington Co., Parry 47, 1874 (NY!;ISC!). Haplopappus cervinus Wats. Amer. Naturalist 7: 301. 1873. Asteraceae. Utah, Antelope Canvon, Wheeler sn, 1872 (US!). Haplopappus nuttallii T. & G. var. depressus Ma- guire .Amer. Midi. Naturalist 37: 144. 1947. Asteraceae. = M. griudelioides var. depressa (Maguire) Cronq. & Keck Millard Co., Desert Experimental Range, .Maguire 20859, 1941 (US!;NY!;UTC!). Haplopappus parryi Gray var. minor Gray Svn. Fl. N. Amer. 1(2): 131. 1884. Asteraceae. = Solidagv parryi (Gray) Greene Salt Lake Co., Alta, Bald Mtn., Jones sn?, 1879(POM!:BRY!;NDG!). Haplopappus scopulorum (Jones) Blake in Tidestr. var. hirtellus Blake Proc. Biol. Soc. Washington 48: 170. 1935. .Asteraceae. Iron Co., Cedar Canyon, Garrett 6051, 1931 (US!;UT!). Hedysarum gremiale Rollins Rhodora 42: 230. 1940. Fabaceae. = H. boreale var. gremiale (Rollins) North- strom & Welsh Uintah Co., 14 mi n Vernal, Rollins 17.33, 19.37 (US!;CAS!;RM!;UTC!). Hedysarum occidentale Greene var. canone Welsh Great Basin Naturalist 38: 314. 1978. Fabaceae. Carbon Co., Soldier Creek, Welsh & Tavlor. 1.52.56, 1977 (BRY!;NY!). Hedysarum utahense Rydb. Bull. Torrey Bot. Club .34: 425. 1907. Fabaceae. = H. boreale var. boreale Salt Lake Co., Salt Lake City, Leonard .55, 1883 (NY!). Helianthella multicaulis D.C. Eaton in Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 170. 1871. Aste- raceae. = H. uniflora (Nutt.) T. & G. Summit Co., Par- leys Park, Watson 605, 1869 (US!;NY!). Helianthus anomalus Blake J. Washington Acad. Sci. 21: .3.33. 1931. Asteraceae. Wayne Co.^ s Hanksville, Stanton 4806, 19,30 (US!;UT!). Helianthus bracteatus E.E. Watson Papers .Michigan Acad. Sci. 9: .393. 1929. Asteraceae. = H. nuttallii T. & G. Cache Co., Logan, Mulford 177, 1898 (MO). June 1982 Welsh: Utah Plant Types 173 Helianthus deserticolus Heiser Proc. Indiana Acad. Sci. 70: 209. 1961. Asteraceae. Washington Co., w Hur- ricane, Stoiitamire 2574, 1957 (IND). Helianthus giganteus var. utahensis D.C. Eaton in Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 169. 1871. Asteraceae. = H. nuttallii T. & G. Wasatch Mts., Watson 603, 1869 (US!). Hennidium alipes Wats. var. pallidum C.L. Porter Rhodora 54: 158. 1952. Nyctaginaceae. = Mirabilis alipes (Wats.) Pi!z Uintah Co., 5 mi sw Vernal, Porter 5308, 1950(RM!;CAS!). Hesperanthes alhomarginata Jones Zoe 2: 251. 1891. Liliaceae. = Eremocrinum albomarginatum (Jones) Jones Emery Co., Green River, Jones sn, 1890 (?). Heuchera rubescens Torr. in Stansb. Explor. Great Salt Lake 388. 1852. Saxifragaceae. Syn: H. versicolor Greene; H. versicolor i. puinila Rosend., Butters, & La- kela Davis (?) Co., Great Salt Lake, Stansbury sn, 1850 (NY!). Heuchera utahensis Rydb. N. Amer. Fl. 22: 114. 1905. Saxifragaceae. = H. parvifolia Nutt. Salt Lake Co., City Creek Canyon, Jones 1458, 1880 (US!;CAS!; RM!;POM!;UT!;BRY!;UTC!). Heuchera versicolor Greene f. pumila Rosend., flut- ters, & Lakela Minnesota Studies PI. Sci. 2: 85. 1936. Sa- xifragaceae. = H. rubescens Torr. Utali? (?). Hieracium utahense Gandg. Bull. Soc. Bot. France 65: 49. 1918. Asteraceae. = H. gracile Hook. Cache Co., Linford sn, 1897 (?). Holodiscus microphyllus Rydb. Bull. Torrey Bot. Club 31: 559. 1904. Rosaceae. = H. dumosus (Nutt.) Heller Salt Lake Co., Alta, Jones 1142, 1879 (P0M!;BRY!;NY!;UTC!). Homalobus canovirens Rydb. Fl. Rocky Mts. ed. 2. 1126. 1922. Fabaceae. = Astragalus coltonii var. moa- bensis Jones Grand Co., La Sal Mts, Rydberg & Garrett 85.36, 1913 (NY!). Homalobus humilis Rydb. Bull. Torrey Bot. Club 34: 417. 1907. Fabaceae. = Astragalus miser var. oblongi- folius (Rydb.) Cronq. Piute Co., n Bullion Creek, Ryd- berg & Carlton 7147, 1905 (NY!;US!;RM!;BRY!). Homalobus paucijugus Rydb. Bull. Torrey Bot. Club 34: 418. 1907. Fabaceae. = Astragalus miser var. tenui- folius (Nutt.) Barneby Syn: A. garrettii Macbr. Salt Lake Co., Big Cottonwood Canyon, Garrett 1580, 1905 (NY!;US!;UT!). Hordeum pusillum Nutt. var. pubens A.S. Hitchc. J. Washington Acad. Sci. 23: 453. 1933. Poaceae. = H. pu- sillum Nutt. Washington Co., LaVerkin, Jones 5196ro, 1894 (POM!). Hosackia rigida Benth. var. nummularia Jones Proc. Calif. Acad. II, 5: 633. 1895. Fabaceae. = Lotus long- ebracteatus Rydb. Washington Co., Rockville, Jones 5224, 1894 (US!;RM!). Houstonia saxicola Eastw. Proc. Calif. Acad. II, 6: 291. 1896. Rubiaceae. = H. rubra Cav. San Juan Co., Butler Spring, Eastwood 33, 1895 (CAS!). Hymenopappus eriopodus A. Nels. Bot. Gaz. 37: 274. 1904. Asteraceae. = H. filifolius var. eriopodus (A. Nels.) Turner Washington Co., Diamond Valley, Goodd- ing 880, 1902 (RM!;US!;NY!). Hymenopappus niveus Rydb. N. Amer. Fl. .34: 52. 1914. Asteraceae. = H. filifolius var. tomentosus (Rydb.) Turner Washington Co., Springdale, Jones 5261, 1894 (US!;NY!). Hymenopappus nudipes Maguire Amer. Midi. Natu- ralist 37: 143. 1947. Asteraceae. = H. filifolius var. al- pestris (Maguire) Shinners Kane Co., 15 mi n Orderville, Maguire 18740, 1940 (NY!;US!). Hymenopappus nudipes Maguire var. alpestris Ma- guire Amer. Mid!. Naturalist 37: 144. 1947. Asteraceae. = H. filifolius var. alpestris (Maguire) Shinners Iron Co., Cedar Breaks, Maguire 19023, 1940 (US!; NY!;UTC!). Hymenopappus pauciflorus Johnst. Contr. Gray Herb. II, 68: 97. 1923. Asteraceae. = H. filifolius var. pauciflorus (Johnst.) Turner San Juan Co., near Bluff, Rydberg & Garrett 9951, I91I (UT!;NY!). Hymenopappus tomentosus Rydb. Bull. Torrey Bot. Club 27: 633. 1900. Asteraceae. = H. filifolius var. to- mentosus (Rydb.) Turner Washington Co., St. George, Palmer 270, 1877 (NY!;ISC!;BRY!). Hymenoxys lemmonii (Greene) Cockerell ssp. greenei Cockerell Bull. Torrey Bot. Club 31: 479. 1904. Aste- raceae. = H. lemmonii (Greene) Cockerell Washington (?) Co., Rock Creek, Palmer 261, 1877 (US!;NY!). Hymenoxys richardsonii (Hook.) Cockerell var. uta- hensis Cockerell Bull. Torrey Bot. Club 31: 477. 1904. Asteraceae. = H. richardsonii (Hook.) Cockerell Emery Co., Emery, Jones 5442, 1894 (US!;POM!). Ivesia utahensis Wats. Proc. Amer. Acad. 10: 71. 1874. Rosaceae. Salt Lake Co., Bald Mt., Jones 1231, 1879(US!;NY!;UTC!)). Juncus canadensis var. kuntzei Buch. Bot. Jahrb. 12: 272. 1890. Juncaceae. = /. tweedyi Rydb. Box Elder Co., near Corinne, Kuntze 3133, 1874 (NY!). Juncus jonesii Rydb. Fl. Rocky Mts. 153, 1061. 1917. Juncaceae. = /. regelii Buch. Salt Lake Co., Alta, Jones 119, 1879(NY!;POM!). Juncus tracyi Rydb. Fl. Rocky Mts. 155, 1061. 1917. Juncaceae. = /. ensifolius Wikstr. Weber Co., Ogden, Tracy 389, 1887 (US!;NY!). Juncus utahensis Martin Rhodora 40: 69. 1938. Jun- caceae. = /. ensifolius var. brunnescens (Rydb.) Cronq. Summit (?) Co., Ashley National Forest, Nord & Sargent 1, 1927 (US!). Juniperus californica var. utahensis Vasey Cat. For- est Trees U.S. 37. 1876. Cuprcssaceae. = /. osteosperma (Torr.) Little Wasatch Mts., Ward sn, 1875 (ISC!). Juniperus californica var. utahensis Engelm. Trans. Acad. Sci. St. Louis 3: 588. 1878. Cupressaceae. = /. os- teosperma (Torr.) Little Syn: /. utahensis (Engelm.) Lem- mon Washington Co., St. George, Palmer? sn, 1877 (?). Kochia americana Wats. var. vestita Wats. Proc. Amer. Acad. 9: 93. 1874. Chenopodiaceae. = K. ameri- cana Wats. Tooele Co., "Tuilla Valley" Watson 991, 1869 (US!;NY!). Krynitzkia echinoides Jones Proc. Calif. Acad. II, 5: 709. 1895. Boraginaceae. = Cryptantha ftdvocanescens var. echinoides (Jones) Higgins Garfield (?) Co., Pahria Canyon, Jones 5297p, 1894 (US!;POM!). Krynitzkia glomerata var. acuta Jones Zoe 2: 250. 1891. Boraginaceae. = Cryptantha wetherillii (Eastw.) Payson Grand Co., Cisco, Jones sn, 1890 (?). Krynitzkia glomerata var. virginensis Jones Contr. W. Bot. 13: 5. 1910. Boraginaceae. = Cryptantha virgi- nensis (Jones) Payson Washington Co., LaVerkin, Jones 5195a, 1894 (US!;RM!;POM!). 174 Great Basin Naturalist Vol. 42, No. 2 Krynitzkia leucophaea var. alata Jones Proc. Calif. Acad. II, 5; 710. 1895. Boraginaceae. = Cryptuntha con- fertiflora (Greene) Payson Kane Co., Johnson, Jones 5289f, 1894 (POM!;US!;'bRY!;NY!). Krynitzkia mensana Jones Contr. W. Hot. 13: 4. 1910. Boraginaceae. = Cryptuntha mensana (Jones) Pay- son Emery Co., Emery, Jones 5445p, 1894 (POM!; US!;RM!). ' Krynitzkia multicaulis var. setosa Jones Cotr. W. Hot. 13: 4. 1910. Boraginaceae. = Cryptantha cinerea (Greene) Cronq. Syn: C. jainexii var. setosa (Jones) Johnst. Millard Co., near Cove Fort, Jones sn, 1901 (POM!). Krynitzkia titahensis Gray Syn Fl. N. Anier. ed.2. 2(1): 427. 1886. Boraginaceae. = Cryptantha iitahensis (Gray) Greene Washington Co., St. George, Palmer 352, 1877 (US!). Krynitzkia watsonii Gray Proc. .Amer. Acad. 20: 271. 1885. Boraginaceae. = Cryptantlia watsonii (Gray) Greene Wasatch Mts., Watson 858, 1869 (US!;NY!). Langloisia setosissima (T. & G.) Greene var. camp- yloclados Brand Pflanzenreich 4. Fam. 250: 171. 1907. Polemoniaceae. = L. setosissima (T. & G.) Greene Washington Co., near St. George, Parry 190, 1874 (US!;ISC!). Laphamia palmeri Gray Proc. Amer. Acad. 13: 372. 1878. Asteraceae. = Perityle tenella (Jones) Macbr. Washington Co., (Note: At Beaverdam in Ariz.) Palmer 199, 1877 (US!;NY!). Laphamia palmeri Gray var. tenella Jones Proc. Ca- lif. Acad. II, 5: 703. 1895. Asteraceae. = Perityle tenella (Jones) Macbr. Washington Co., Springdale, Jones 52491, 1894 (US!;POM!;NY!;BRY!). Laphamia stansburyi Gray PI. Wright. 1: 101. 1852. Asteraceae. = Perityle stansburyi (Gray) Macbr. Tooele Co., Stansbury Island, Stansbury sn, 1850 (NY!;BRY!). Lappula collina Greene Pittonia 4: 96. 1899. Boragi- naceae. = L. oeeidentalis (Wats.) Greene sens. lat. Piute Co., Kingston, Jones sn, 1894 (:'). Lathyrus brachycalyx Rydb. var. brachycalyx Bull. Torrev Bot. Club 34: 425. 1907. Fabaceae. Salt Lake Co., City Creek Canyon, Leonard 101, 1883 (NY!). Lathyrus coriaceus White Bull. Torrey Bot. Club 21: 452. 1894. Fabaceae. = L. lanzwertii var. lanzwertii Wasatch Mts., Watson 297, 1869 (US!). Lathyrus utahensis Jones Proc. Calif. Acad. II, 5: 678. 1895. Fabaceae. = L. puuciflorus var. utahensis (Jones) Peck Sevier Co., Irelands Ranch, Jones 54411, 1894 (US!;NY!). Lathyrus zionis C.L. Hitchc. Univ. Washington Publ. Biol. 15: 36. 1952. Fabaceae. = L. braehyealyx var. zi- onii (C.L. Hitchc.) Welsh Kane Co., 10 mi e Zion N.P., Hitchcock 19013, 1949(CAS!,RM!;UTC!;NY!). Lepidium brachybotryum Rydb. Bull. Torrey Bot.club 34: 427. 1907. Brassicaceae. = L. montanum var. montanum Juab Co., Juab, Goodding 1075, 1902 (NY!;US!;RM!). Lepidium georginum Rydb. Bull. Torrey Bot. Club 30; 253. 1903. Brassicaceae. = L. lasiocarpum var. georgi- num (Rydb.) C.L. Hitchc. Washington Co., Southern Utah, Parry 19, 1874 (NY!;ISC!). Lepidium jonesii Rydb. Bull. Torrey Bot. Club 29; 233. 1902. Brassicaceae. = L. montanum var. jonesii (Rydb.) C.L. Hitchc. Washington Co., St. George, Jones 1636, 1880 (US!;CAS!;RM!;POM!;NY!). Lepidium montanum Nutt. in T. & G. var. alpinum Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 29. 1871. Brassicaceae. Salt Lake Co., Cottonwood Canyon, Watson 122, 1869 (US!;NY!). Lepidium montanum Nutt. in T. & G. var. demissum C.L. Hitchc. Madrofio 10: 157. 1950. Brassicaceae. = L. barnebyanum Reveal Duchesne Co., sw Duchesne, Rip- ley & Barneby 8699, 1947 (US!;CAS!;NY!). Lepidium montanum Nutt. in T. & G. var. neeseae Welsh & Reveal Great Basin Naturalist 34: 334. 1977. Brassicaceae. Garfield Co., Hells Backbone, Neese & White 3332, 1977 (BRY!;NY!;UT!). Lepidium montanum Nutt. in T. & G. var. stellae Welsh & Reveal Great Basin Naturalist 34: .3.34. 1977. Brassicaceae. Kane Co., se Cannonville, Welsh & Welsh 12841, 1975 (BRY!). Lepidium ostler Welsh & Goodrich Great Basin Natu- ralist 40; 80. 1980. Brassicaceae. Beaver Co., Frisco, Os- tler & Anderson 1258, 1978 (BRY!). Lepidium utahense Jones Zoe 4: 266. 1893. Brassi- caceae. = L. integrifolium Nutt. Beaver Co., Milford, Jones 1821, 1880 (POM!;NY!;BRY!;UT!;UTC!;ISC!). Lepidium zionis A. Nels. Bot. Gaz. 42: 50. 1906. Brassicaceae. = L. integrifolium Nutt. Sevier Co., Rich- field, Jones 5411, 1894 (RM!;BRY!;POM!;NY!). Leptodactylon brevifolium Rydb. Bull Torrey Bot. Club. 40: 474. 1913. Polemoniaceae. = L. pungens (Torr.) Nutt. Iron Co., Juniper Range, Purpus 6306, 1898 (US!). Leptotaenia eatonii Coult. & Rose Rev. N. Amer. Umbell. 52. 1888. Apiaceae. = Lomatium disseetum var. eatonii (Coult. & Rose) Cronq. Utah, Eaton 147, 1869 (?). Lesquerella garrettii Payson Ann. Missouri Bot. Card. 8; 213. 1921. Brassicaceae. Salt Lake Co., Big Cotton- wood Canyon, Garrett 1.344, 1908 (MO!;RM!). Lesquerella gordonii var. sessilis Wats. Proc. Amer. Acad. 23; 253. 1888. Brassicaceae. = L. tenella A. Nels. Washington Co., near St. George, Parry sn, 1874 (GH). Lesquerella hemiphysaria Maguire var. hemiphysaria .\mer. Midi. Naturalist 27: 456. 1942. Brassicaceae. San- pete Co., Wasatch Plateau, Maguire 20053, 1940 (UTC!). Lesquerella hemiphysaria Maguire var. lucens Welsh & Reveal Great Basin Naturalist 37: .338. 1977. Bra.ssi- caceae. Carbon Co., Range Creek, Welsh & Taylor 151.39, 1977 (BRY!). Lesquerella hitchcockii ssp. tumulosa Barneby Leafl. W. Bot. 10: 313. 1966. Brassicaceae. = L. tumulosa (Barnebv) Reveal Kane Co., se Cannonville, Barneby 14424, 1966(US!;BRY!;UTC!;NY!). Lesquerella multiceps Maguire Amer. Midi. Natural- ist 27: 465. 1942. Brassicaceae. Cache Co., Tony Grove Lake, Maguire 16030, 1938 (UTC!;RM!). Lesquerella rubicundula Rollins Contr. Dudley Herb. 3: 178. 1941. Brassicaceae. Garfield Co., Red Canyon, Egglcston 8198, 1912 (US!). Lesquerella subumbellata Rollins Rhodora 57: 255. 1955. Brassicaceae. Uintah Co., n Vernal, Rollins 17,5, 19.37 (US!;RM!;NY!;UTC!). Lesquerella utahensis Rydb. Bull. Torrey Bot. Club .30: 252. 1903. Brassicaceae. Utah Co., American Fork Canyon, Jones 1.354, 1880 (NY!;US!;CAS!). Lesquerella wardii Wats. Proc. Amer. Acad. 23; 252. 1888. Brassicaceae. Garfield Co., Aquarius Plateau, Ward 589, 1875 (GH;US!;RM!). June 1982 Welsh: Utah Plant Types 175 Ligusticum brevilobum Rydb. Fl. Rocky Mts. 613, 1064. 1917. Apiaceae. = L. porteri Coult. & Rose Gar- field Co., Aquarius Plateau, Rydberg & Carlton 7473, 1905 (NY!;US!). Ligusticum filicimim Wats. Proc. Anier. Acad. 11: 140. 1876. Apiaceae. Summit (?) Co., Uinta Mts., Wat- son 454, 1869 (US!;NY!). Limnia utahensis Rydb. Bull. Torrey Bot. Club 39: 314. 1912. Portulacaceae. = Montia perfoliata (Donn) Howell Washington Co., St. George, Palmer 56, 1877 (NY!). Linosyris serrulata Torr. in Stansb. Explor. Great Salt Lake 389. 1852. Asteraceae. = Chrijsothamnus vis- cidiflorus var. viscidiflorus Salt Lake Co., Salt Lake Val- ley," Stansbury sn. 1850 (NY!). Linum aristatum Engelm. in Wisliz. var. subteres Trel. in Eastwood Proc. Calif. Acad. II, 6: 285. 1896. Li- naceae. = L. subteres (Trel.) Winkler San Juan Co., Willow Creek, Eastwood sn, 1895 (?). Linum kingii Wats. Rep. U.S. Geol. Explor. 40th Par- allel, Bot. 5: 49. 1871. Linaceae. Summit Co., Uintas, Watson 203, 1869 (US!). Linum kingii Wats. var. pinetorum Jones Proc. Calif. Acad. II, 5: 628. 1895. Linaceae. = L. kingii Wats. Gar- field Co., canyon above Tropic, Jones 5306, 1894 (US!;F0M!;NY!;'BRY!). Lomatium jonesii Coult. & Rose Contr. U.S. Natl. Herb. 7: 233. 1900. Apiaceae. = L. foeniculaceum var. macdougalii (Coult. & Rose) Cronq. Sevier Co., Irelands Ranch, Jones 5435, 1894 (US!). Lomatium junceum Barneby & Holmgren Brittonia 31: 96. 1979. Apiaceae. Emery Co., San Rafael Swell, Holmgren et al. 8778, 1978 (NY!;BRY!;UTC!). Lonicera utahensis Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 133. 1871. Caprifoliaceae. Salt Lake Co., Cottonwood Canyon, Watson 477, 1869 (US!;NY!). Lotus longebracteatus Rydb. Bull. Torrey Bot. Club .30: 254. 1903. Fabaceae. Syn: L. oroboides var. niimmii- Jarius (Jones) Isely Washington Co., "South Utah" ("Mo- kaik Pass"), Palmer 94, 1877 (US!;NY!;BRY!). Lotus utahensis Ottley Brittonia 5: 108. 1944. Fa- baceae. Kane Co., w Carmel, Ferguson & Ottley 5613 (?)■ Lupinus aegraovium C.P. Sm. Sp. Lupinorum 694. 1951. Fabaceae. = L. sericeus var. sericeus sens. lat. Se- vier Co., Salina Experiment Station, Huffman 49-9, 1949 (CAS!). Lupinus aridus var. utahensis Wats. Proc. Amer. Acad. 8: 5.34. 1873. Fabaceae. = L. caespitosus Nutt.; Svn; L. watsonii Heller Summit Co., Parleys Park, Wat- son 2.34, 1869 (US!). Lupinus barbiger Wats. Proc. Amer. Acad. 8: .528. 1873. Fabaceae. = L. sericeus var. barbiger (Wats.) Welsh Kane Co., S. Utah, Siler sn, 1872? (GH). Lupinus eatonanus C.P. Sm. Sp. Lupinorum 671. 1949. Fabaceae. = L. leucophyllus Dougl. Wasatch Co., Hailstone, Smith .3995, 1925 (CAS!). Lupinus garrettianus C.P. Sm. Sp. Lupinorum 672. 1949. Fabaceae. = L. argenteus var. argenteus Duch- esne Co., 1 mi w Duchesne, Garrett 8.303, 1940 (UT!). Lupinus holosericeus var. utahensis Wats. Proc. Amer. Acad. 8: 5.33. 1873. Fabaceae. = L. caudatus var. caudatus Salt Lake Co. ?, Watson sn?, 1869 (US!). Lupinus huffmanii C.P. Sm. Sp. Lupinorum 693. 1951. Fabaceae. = L. sericeus var. sericeus sens. lat. Se- vier Co., Salina Experiment Station, Huffman 47-7A, 1949 (CAS!). Lupinus jonesii Rydb. Bull. Torrey Bot. Club 30: 256. 1903. Fabaceae. Syn: L. leucanthus Rydb. Washington Co., Silver Reef, Jones 514.3, 1894 (US!;RM!;BRY!;NY!). Lupinus larsonanus C.P. Sm. Sp. Lupinorum 696. 1951. Fabaceae. = L. sericeus var. sericeus sens. lat. Se- vier Co., inter Browns Hole & Hoodoo, Larson sn, 1934 (CAS!). Lupinus leucanthus Rydb. Bull. Torrey Bot. Club .30; 259. 1903. Fabaceae. = L. jonesii Rydb. Washington Co., Springdale, Jones .5249e, 1894 (US!;RM!; BRY!;POM!;NY!). Lupinus leucophyllus Dougl. ex Lindl. var. lupinus Rydb. Bull. Torrey Bot. Club 40: 44. 1913. Fabaceae. Syn: L. utahensis Moldenke = L. caudatus var. caud- atus San Juan Co., Bears Ears, Rydberg &c Garrett 9363, 1911 (NY!). Lupinus maculatus Rydb. Bull. Torrey Bot. Club 30: 257. 1903. Fabaceae. Utah Co., Pleasant Valley (P.V.) Jet., Jones sn, 1883 (NY!;US!;POM!;UTC!). Lupinus marianus Rydb. Bull. Torrey Bot. Club 34: 41. 1907. Fabaceae. = L. sericeus var. marianus (Rydb.) Welsh Piute Co., Bullion Creek, Rydberg & Carlon 7024, 1905(US!;RM!;NY!). Lupinus prunophilus Jones Contr. W. Bot. 13: 7. 1910. Fabaceae. Juab Co., Robinson, Jones sn, 1909 (CAS!;POM!;BRY!;NY!). Lupinus pulcher Eastw. Leafl. W. Bot. 3: 173. 1942. Fabaceae. = L. hillii Greene Iron Co., 18 mi s Cedar City, Barkley & Reed 4069, 1939 (CAS!). Lupinus puroviridis C.P. Sm. Sp. Lupinorum 694. 1951. Fabaceae. = L. sericeus var. sericeus sens. lat. Se- vier Co., Salina Experiment Station, Huffman 49-7B, 1949 (CAS!). Lupinus quercus-jugi C.P. Sm. Sp. Lupinorum 696. 1951. Fabaceae. = L. sericeus var. sericeus sens. lat. Se- vier Co., Browns Hole, Clawson C241, 1935 (CAS!). Lupinus rickeri C.P. Sm. Sp. Lupinorum 695. 1951. Fabaceae. = L. sericeus var. sericeus sens. lat. Sevier Co., Salina Experiment Station, Ricker 14080, 1917 (CAS!). Lupinus rubens Rydb. Bull. Torrey Bot. Club 34: 45. 1907. Fabaceae. = L. pusiUus var. rubens (Rydb.) Welsh Washington Co., St. George, Parry 41, 1874 (NY!;ISC!). Lupinus salinensis C.P. Sm. Sp. Lupinorum 695. 1951. Fabaceae. = L. sericeus var. sericeus sens. lat. Se- vier Co., Salina Experiment Station, Clawson C.202, 1935 (CAS!). Lupinus sileri Wats. Proc. Amer. Acad. 10: 345. 1875. Fabaceae. = L. kingii Wats. Southern Utah, Siler sn (GH). Lupinus spatulatus Rydb. Bull. Torrey Bot. Club 29: 244. 1902. Fabaceae. = L. argenteus var. boreus (C.P. Sm.) Welsh Wasatch Mts., Watson 225, 1869 (NY!). Lupinus tooelensis C.P. Sm. Sp. Lupinorum 640. 1948. Fabaceae. = L. prunophilus Jones Tooele Co., Deep Creek Mts., Cottam 7203, 1937 (UT!). Lupinus watsonii Heller Muhlenbergia 1: 114. 1905. Fabaceae. = L. caespitosus Nutt. Summit Co., Parleys Park, Watson 234, 1869 (US!). 176 Great Basin Naturalist Vol. 42, No. 2 Lychnis kingii Wats. Proc. Amer. Acad. 12: 247. 1877. Caryophyllaceae. Summit Co., head Bear River, Watson 153,1869 (US!;NY!). Lygodesmia entrada Welsh & Goodrich Great Basin NaturaHst 40: 83. 1980. Asteraceae. Grand Co., Tusher Canyon, Welsh & Welsh 16725, 1978 (BRY!;UT!;NY!). Lygodesmia grandiflora (Nutt.) T. & G. var. stricta Maguire Amer. Midi. Naturalist 37: 145. 1947. Aste- raceae. Carbon Co., 1 mi s Price, Maguire 18417, 1940 (NY!;US!;UTC!). Lygodesmia juncea (Pursh) D. Don var. dianthopsis D.C. Eaton Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 200. 1871. Asteraceae. = L. dianthopsis (D.C. Eaton) Tomb Davis (?) Co., Great Salt Lake Islands, Watson 707, 1869 (US!). Machaeranthera commixta Greene Pittonia 4: 71. 1899. Asteraceae. Syn: M. leptophylla Rydb.? Garfield Co., Bromide Pass, Jones 5695v, 1894 (US!). Machaeranthera glabriuscula (Nutt.) Cronq. & Keck var. confertiflora Cronq. Leafl. W. Bot. 10: 11. 1963. Asteraceae. = Xylorhiza confertiflora (Cronq.) T.J. Wat- son Garfield Co., 11 mi ne Henrieville, Cronquist 9164, 1961 (NY!;UTC!). Machaeranthera kingii (D.C. Eaton) Cronq. & Keck var. bamebyana Welsh & Goodrich in Welsh Brittonia .33: 299. 1981. Asteraceae. Millard Co., Canyon Mts., Goodrich 14929, 1980 (BRY!;GH!;NY!;US!;RM!; UC!;USFS!;UT!;UTC!). Machaeranthera latifolia A. Nels. Proc. Biol. Soc. Washington 20: .38. 1907. Asteraceae. Syn: M. rubr- icaulis Rydb.; Aster nihrotinctiis Blake = M. bigelovii Gray ? Salt Lake Co., Big Cottonwood Canyon, Garrett 1933, 1906(US!;RM!;UT!). Machaeranthera leptophylla Rydb. Bull. Torrey Bot. Club 37: 147. 1910 Asteraceae. = M. commixta Greene? Cache Co., Logan, Rydberg sn, 1895 (NY!;BRY!). Machaeranthera paniculata A. Nels. Proc. Biol. Soc. Washington 20: 38. 1907. Asteraceae. = M. bigelovii Gray ? Salt Lake Co., Parleys Canyon, Garrett 2083, 1906(US!;RM!;UT!). Machaeranthera pulverulenta var. vacans A. Nels. Bot. Gaz. 56: 70. 1913. Asteraceae. = M. canescens (Pursh) Gray? San Juan Co., Walker 360, 1912 (RM!;NY!). Machaeranthera tortifolia (T. & G.) Cronq. & Keck var. imberbis Cronq. Leafl. W. Bot. 10: 12. 1963. Aste- raceae. = Xylorhiza tortifolia var. iinberbis (Cronq.) T.J. Watson Grand Co., w Moab, Cronquist 8994, 1961 (NY!;UTC!). Macronema obovatum Rydb. Bull. Torrey Bot. Club 27: 618. 1900. Asteraceae. = Haplopappus rydbergii Blake Salt Lake Co., City Creek Canyon, Jones 1081, 1895(US!;CAS!;RM!;POM!;NY!;UTC!). Madronella oblongifolia Rydb. Bull. Torrey Bot. Club 36: 686. 1909. Lamiaceae. = Monardella odoratissima Benth. Juab Co., Mt. Nebo, Rydberg & Carlton 7706, 1905(NY!;US!;RM!). Madronella sessilifolia Rydb. Bull. Torrey Bot. Club 36: 685. 1909. Lamiaceae. = Monardella odoratissima Benth. Washington Co., St. George, Palmer .393, 1877 (NY!). Mammilluria chlorantha Engelm. in Wheeler Rep. U.S. Geogr. Surv. W. 100th Meridian 6: 127. 1878. Cac- taceae. = Coryphantha vivipara var. deserti (Engelm.) W.T. Marshall Washington Co., e St. George, Parry sn, 1874 (ISC!;BRY!). Mentzelia albicaulis Dougl. in Hook. var. integrifolia Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 114. 1871. Loasaceae. = M. dispersa Wats. Davis Co., Ante- lope Island, Watson 4.30, 1869 (US!;NY!). Mentzelia argillosa Darlington Ann. Missouri Bot. Card. 21: 153. 1934. Loasaceae. Sevier Co., Vermillion, Jones 5631, 1894 (MO;POM!;US!). Mentzelia multiflora (Nutt.) Gray var. integra Jones Proc. Calif. Acad. Sci 11, 5: 689. 1895. Loasaceae. = M. integra (Jones) Tidestr. Washington Co., Rockville, Jones 6082C, 1894 (US!). Mentzelia pterosperma Eastw. Proc. Calif. Acad. 11, 6: 290. 1896. Loasaceae. San Juan Co., Willow Creek, East- wood 31, 1895 (CAS!). Mertensia brevistyla Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 239. 1871. Boraginaceae. Summit (?) Co., Wasatch Mts., Watson 845, 1869 (NY!). Mertensia leonardii Rydb. Bull. Torrey Bot. Club 36: 681. 1909. Boraginaceae. = M. arizonica Greene var. leonardii (Rydb.) Johnst. Salt Lake Co., Mill Creek Can- yon, Leonard sn, 1884 (NY!). Mertensia paniculata var. nivalis Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 239. 1871. Boragi- naceae. = M. viridis A. Nels. var. cana (Rydb.) L.O. Williams Summit Co., Bear River Canon, Watson 844, 1869 (CAS!;US!;NY!). Mertensia praecox Smiley in Macbr. Contr. Gray Herb. 11, 48: 10. 1916. Boraginaceae. = M. oblongifolia var. nevadensis (A. Nels.) L.O. Williams Cache Co., Lo- gan Canyon, Smith 2160, 1910 (UTC!). Mertensia sampsonii Tidestr. Proc. Biol Soc. Wash- ington 26: 122. 1913. Boraginaceae. = M. arizonica var. leonardii (Rydb.) Johnst. Sanpete Co., e of Ephraim, Sampson 677, (US!). Mertensia toyabensis var. subnuda Macbr. Contr. Gray Herb. II, 48: 7. 1916. Boraginaceae. = M. arizona var. subnuda (Macbr.) L.O. Williams Wayne Co. (?), Fish Lake Mountain. Ward 329, 1875 (GH). Micropuntia barkleyana Daston Amer. Midi. Natu- ralist 36: 662. 1946. Cactaceae. = Opuntia pulchella Engelm. Millard Co., Desert Experimental Range, Marsh sn, 1945 (F?). Micropuntia brachyrohopalica Daston Amer. Midi. Naturalist 36: 661. 1946. Cactaceae. = Opuntia pul- chella Engelm. Millard Co., Desert Experimental Range, Marsh sn, 1945 (F?). Micropuntia spectatissima Daston Amer. Midi. Natu- ralist 36: 661. 1946. Cactaceae. = Opuntia pulchella Engelm. Millard Co., Desert Experimental Range, Marsh sn, 1946 (F?). Mimulus eastwoodiae Rydb. Bull. Torrey Bot. Club 40: 483. 1913. Scrophulariaceae. San Juan Co., San Juan River, near Bluff, Rydberg 9883, 1911 (NY!; US!;RM!;UT!). Mimulus glabratus H.B.K. ssp. utahensis Pennell Acad. Nat. Sci. Philadelphia Monogr. 1: 123. 1935. Scro- phulariaceae. = M. glabratus H.B.K. Millard Co., Preuss Lake, Tidestrom 11180, 1919 (PH). June 1982 Welsh: Utah Plant Types 177 Mimulus parryi Gray Proc. Amer. Acad. 11: 97. 1876. Scrophulariaceae. Washington Co., near St. George, Par- ry 147, 1874 (NY!;US!;ISChNDG!). Mirabilis gltitinosa A. Nels. Proc. Biol. Soc. Washing- ton 17: 92. 1904. Nyctaginaceae. = M. bigelovii Gray var. retrorsa (Heller) Munz Washington Co., near St. George, Goodding 778. 1902 (RM!). Mitella stenopetala Piper Erythea 7: 161. 1899. Sa- xifragaceae. = M. stauropetala Piper Salt Lake (?) Co., Wasatch Mts., Watson 365, 1869 (?). Muhlenbergia ctirtifolia Scribn. Bull. Torrey Bot. Club 38: 328. 1911. Poaceae. = M. thurberi Rydb. Kane Co., inter Kanab & Carmel, Jones 6077j, 1894 (US!). Najas flexilis ssp. caespitosus Maguire in Maguire & Jensen Rhodora 44: 7. 1942. Najadaceae. = N. caespi- tosus (Maguire) Reveal Sevier Co., Pelican Point, Fish Lake, Maguire 19888, 1940 (UTC!;BRY!). Nasturtium obtusum Nutt. var. alpinum Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 15. 1871. Brassi- caceae. = Rorippa curvipes var. alpina (Wats.) Stuckey Summit Co., head of Bear River Canyon, Uinta Mts., Watson 60, 1869 (US!). Navarretia setosissitna T. & G. in Ives Rep. Colorado Riv. W. 4: 22. 1860. Polemoniaceae. = Langloisia seto- sissima (T. & G.) Greene Washington Co., Virgin River, Fremont 414, (NY!). Nemophila breviflora Gray Proc. Amer. Acad. 10: 315. 1875. Hydrophvllaceae. Summit Co., Parleys Park, Watson 169, 1869 (NY!;US!). NothoJaena parryi D.C. Eaton Amer. Naturalist 9: 351. 1875. Polvpodiaceae. Washington Co., St. George, Parry 263, 1874 (US!;ISC!;BRY!). Nuttallia lobata Rydb. Bull. Torrey Bot. Club 40: 61. 1913. Loasaceae. = Mentzelia Integra (Jones) Tidestr. Washington Co., near St. George, Palmer 172, 1877 (NY!). Oenothera acutissima W.L. Wagner Systematic Bot- any 6: 153. 1981. Onagraceae. Daggett Co., Greendale Campground, Neese & Peterson 5428, 1978 (BRY!). Oenothera albicaulis Pursh var. decumbens Wats, ex Parry Amer. Naturalist 9: 270. 1875. Onagraceae. = O. deltoides ssp. ambigua (Wats.) W. Klein Washington Co., near St. George, Parry 63, 1874 (GH?). Oenothera alyssoides H. & A. var. minutiflora Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 111. 1871. Onagraceae. = Camissonia minor (A. Nels.) Raven Tooele Co.. Stansbury Island, Watson 421, 1869 (US!). Oenothera alyssoides Pursh var. villosa Wats. Proc. Amer. Acad. 8: 591. 1873. Onagraceae. = Camissonia boothii ssp. alyssoides (H. & A.) Raven Salt Lake (?) Co., near Salt Lake, Stansbury sn, 1850 (NY!). Oenothera ambigua Wats. Proc. Amer. Acad. 14: 293. 1879. Onagraceae. = O. deltoides var. ambigua (Wats.) Munz Washington Co., near St. George, Palmer 162, 1877 (US!;ISC!). Oenothera brevipes var. parviflora Wats, in Parry Amer. Naturalist 9: 271. 1875. Onagraceae. = Cam- issonia multijuga (Wats.) Raven Washington Co., near St. George, Parry 74, 1874 (GH;MO;F). Oenothera caespitosa Nutt. var. jonesii Munz Amer. J. Bot. 18: 731. 1931. Onagraceae. Syn: O. caespitosa ssp. jonesii (Munz) Munz Tooele Co., Fish Springs, Jones sn, 1891 (POM!). Oenothera californica ssp. avita W. Klein Aliso 5: 179. 1962. Onagraceae. = O. avita (W. Klein) W. Klein Washington Co., 0.5 ne of Leeds, Klein 1049, 1959 (RSA). Oenothera gauraeflora var. hitchcockii H. Lev. Mon- ogr. Oenothera 226. 1905. Onagraceae. = Camissonia boothii ssp. ah/ssoides (H. & A.) Raven Simpsons Park, without coll., 1859 (MO). Oenothera hookeri T. & G. var. angustifolia Gates Mut. Factors Evol. 10: 30. 1915. Onagraceae. Utah Co., Asphalt (s Thistle), Jones 5624, 1894 (US!;NY!). Oenothera johnsonii Parry Amer. Nat. 9: 270. 1875. Onagraceae. = O. primiveris Gray Washington Co., near St. George, Parry 64, 1874 (ISC!). Oenothera hngissima Rydb. Bull. Torrey Bot. Club 40: 65. 1913. Onagraceae. San Juan Co., Natural Bridges Rydberg & Garrett 9410, 1911 (NY!;US!). Oenothera multijuga Wats. Amer. Naturalist 7: 300. 1873. Onagraceae. = Camissonia multijuga (Wats.) Rav- en Kane Co., near Kanab, Thompson sn, 1872 (NY?). Oenothera multijuga Wats. var. orientalis Munz Amer. J. Bot. 15: 232. 1928. Onagraceae. = Camissonia walkeri ssp. walkeri Grand Co., Moab, Jones sn, 1913 (POM!). Oenothera parryi Wats, ex Parry Amer. Naturalist 9: 270. 1875. Onagraceae. = Camissonia parryi (Wats.) Raven Washington Co., near St. George, Parry 72, 1874 (US!;NY!;BRY!;ISC!;NDG!). Oenothera scapoidea Nutt. in T. & G. ssp. utahensis Raven Univ. Calif. Publ. Bot. 34: 96. 1962. Onagraceae. = Camissonia scapoidea ssp. utahensis (Raven) Raven Salt Lake Co., Black Rock, Watson 414, 1869 (US!). Oenothera tenuissima Jones Proc. Calif. Acad. II, 5: 683. 1895. Onagraceae. = Camissonia parryi (Wats.) Raven Washington Co., Rockville, Jones 6083, 1894 (POM!;US!;RM!;NY!). Oenothera triloba var. ecristata Jones Proc. Calif. Acad. II, 5: 681. 1895. Onagraceae. = O. flava (A. Nels.) Garrett Garfield Co., Panguitch Lake, Jones 60155, 1894 (POM!). Opuntia barhata M. Brandegee ex J. A. Purpus Mon- ats. Kakteenk. 10: 97. 1900. Cactaceae. = O. poly- acantha (?) Haw. San Juan (?) Co., La Sal Mts., Purpus sn, 1897 (?). Opuntia barbata M. Brandegee ex J.A. Purpus var. gracillima M. Brandegee ex J.A. Purpus Monats. Kak- teenk. 10: 110, 120. 1900. Cactaceae. = O. polyacantha (?) Haw. San Juan (?) Co., La Sal Mts., Purpus sn, 1897 (?)• Opuntia basilaris Engelm. & BigeL var. woodburyi Earl Sahuaroland Bull. 34: 15. 1980. Cactaceae. Wash- ington Co., Fort Pierce Wash, Woodbury 2060a, 1977 (BRY!). Opuntia palmeri Engelm. Contr. U.S. Natl. Herb. 3: 423. 1896. Cactaceae. = ? O. phaeacantha var. discata (Griffiths) Benson & Walkington Washington Co., near St. George, Palmer sn, 1877 (MO). Opuntia rubrifolia Engelm. Contr. U.S. Natl. Herb. 3: 424. 1896. Cactaceae. = O. erinacea var. ursina Par- ish Washington Co., near St. George, Palmer 3, 1877 (MO). Opuntia utahensis J.A. Purpus Monats. Kakteenk. 19: 133. 1909. Cactaceae. = O. macrorhiza Engelm. San Juan (?) Co., La Sal Mts., Purpus sn, 1897 (US!). 178 Great Basin Naturalist Vol. 42, No. 2 Oreocarya breviflora Osterh. in Payson Univ. Wyom- ing Publ. Bot. 1: 169. 1926. Boraginaceae. = Crijptantha breviflora (Osterh.) Payson Uintah Co., n Jensen, Os- terhout 6414, 1925 (RM!;BRY!). Oreocarya commixta Macbr. Contr. Gray Herb. II, 48: 33. 1916. Boraginaceae. = Cryptantha humilis var. commixta (Macbr.) Higgins Juab Co., Juab, Goodding 1074, 1902 (GH;RM!). Oreocarya disticha Eastw. Bull. Torrey Bot. Club 30: 258. 1903. Boraginaceae. Syn: Cryptantha jamesii var. disticha (Eastw.) Payson = C. cinerea (Greene) Cronq. San Juan Co., Bartons range, Eastwood sn, 1895 (CAS!;NY!). Oreocarya dolosa Macbr. Contr. Gray Herb. II, 48: 32. 1916. Boraginaceae. = Cryptantha humilis var. shantzii (Tidestr.) Higgins Cache Co., Logan, Smith 1605, 1909 (RM!). Oreocarya jonesiana Payson Univ. Wyoming Publ. Bot. 1: 168. 1926. Boraginaceae. = Cryptantha jone- siana (Payson) Payson Emery Co., San Rafael Swell, Jones sn, 1914 (RM!;POM!;BRY!). Oreocarya pustulosa Rydb. Bull. Torrev Bot. Club 40: 480. 1913. Boraginaceae. = Cryptantha pustulosa (Rydb.) Payson Syn: C. jamesii var. pustulosa (Rydb.) Harrington San Juan Co., Elk Mts., Rydberg & Garrett 9320, 1911 (NY!). Oreocarya rugulosa Payson Univ. Wyoming Publ. Bot. 1: 166. 1926. Boraginaceae. = Cryptantha rugulosa (Payson) Payson Tooele Co., Fish Springs, Jones sn, 1891 (GH;RM!;POM!). Oreocarya shantzii Tidestr. Proc. Biol. Soc. Washing- ton 26: 122. 1913. Boraginaceae. = Cryptantha humilis var. shantzii (Tidestr.) Higgins Salt Lake Co., s Great Salt Lake, Kearney & Shantz 3098, 1912 (US!). Oreocarya tenuis Eastw. Bull. Torrey Bot. Club 30: 244. 1903. Boraginaceae. = Cryptantha tenuis (Eastw.) Payson Grand Co., Courthouse Wash, Eastwood sn, 1892(CAS!;RM!;NY!). Oreocarya torva A. Nels. Amer. J. Bot. 23: 269. 1936. Boraginaceae. = Cryptantha flava (A. Nels.) Payson Carbon Co., Price, Flowers 6395, 1933 (UTl). Oreocarya wetherillii Eastw. Bull. Torrey Bot. Club 30: 242. 1903. Boraginaceae. = Cryptantha wetherillii (Eastw.) Payson Grand Co., Courthouse Wash, East- wood sn, 1892 (CAS!;RM;NY!). Oreocarya iLnlliamsii A. Nels. Amer. J. Bot. 21: 578. 1934. Boraginaceae. = Cryptantha stricta (Osterh.) Pay- son Daggett Co., Flaming Gorge, Williams 489, 1932 (RM!;NY!). Orogenia linearifolia Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 120. 1871. Apiaceae. Summit Co., Wasatch Mts., north of Parleys Park, Watson 440, 1869 (US!;NY!). Oxybaphus angustifolia var. viscidus Eastw. Proc. Calif. Acad. II, 6: 313. 1896. Nyctaginaceae. = O. lin- earis (Pursh) Robins. San Juan Co., Butler Spring, East- wood 101, 1895 (CAS!). Oxybaphus glaber Wats. Amer. Naturalist 7: 301. 1873. Nyctaginaceae. Kane Co., Kanab, Thompson sn, 1872 (US!). Oxytropis jonesii Barneby Proc. Calif. Acad. IV, 27: 215. 1952. Fabaceae. Garfield Co., Red Canyon, Ripley & Barneby 8550, 1947 (CAS!). Oxytropis oreophila Gray var. juniperina Welsh Great Basin Naturalist 38: 339. 1978. Fabaceae. Wayne Co., 1 mi e Bicknell, Welsh & Moore 13828, 1976 (BRY!;UT!;ISC!). Pachylophus crinitis Rydb. Fl. Rocky Mts. 598, 1064. 1917. Onagraceae. = Oenothera caespitosa var. crinita (Rydb.) Munz Wayne Co., Rabbit Valley, Ward 526, 1875 (US!). Paronychia pulvinata Gray var. longiaristata Chaudri Revis. Paronychia 194. 1968. Caryophyllaceae. Duchesne Co., Mt. Emmons, Hermann 5041, 1933 (BRY!). Parrya platycarpa Rydb. Bull. Torrey Bot. Club 39: 326. 1912. Brassicaceae. = P. rydbergii Botsch. Summit (?) Co., Uinta Mts, Watson 54, 1869 (US!;NY!). Parthenium alpinum (Nutt.) T. & G. var. ligulatum Jones Contr. W. Bot. 13: 16. 1910. Asteraceae. Syn: Bolophyta ligulata (Jones) W.A. Weber = P. ligulatum (Jones) Barneby Duchesne Co., Theodore, Jones sn, 1908 (POM!;US!;CAS!;BRY!). Pediocactus despainii Welsh & Goodrich Great Basin Naturalist 40: 83. 1980. Cactaceae. Emery Co., San Ra- fael Swell, Despain 266a, 1978 (BRY!;NY!)' Pediocactus hermannii W.T. Marshall Saguaroland Bull. 8: 78. 1954. Cactaceae. = P. simpsonii (Engelm.) Britt. & Rose Garfield Co., Hatch, Hermann & Hermann sn, 1953 (BRY!). Pediocactus winkleri Heil Cact. &: Succ. J. (U.S.) 51: 28. 1979. Cactaceae. Wayne Co., Winkler sn, (UNM). Penstemon abietinus Pennell Contr. U.S. Natl. Herb. 20: 376. 1920. Scrophulariaceae. Sevier Co., Irelands Ranch, Jones 5440, 1894 (US!;RM!;BRY!;NY!). Penstemon acuminatus Dougl. var. congestus Jones Proc. Calif. Acad. II, 5: 714. 1895. Scrophulariaceae. Syn: P. pachyphyllus ssp. congestus (Jones) Keck; do var. congestus (Jones) N. Holmgr. = P. congestus (Jones) Pennell Washington Co., near Cannan Ranch, Rockwell, Jones 5262, 1894 (POM!;US!). Penstemon angustifolius Nutt. var. vernalensis N. Holmgren Brittonia 31: 229. 1978. Scrophulariaceae. Uintah Co., n Maeser, Holmgren et al. 8748, 1979 (NY!;BRY!;UTC!). Penstemon atwoodii Welsh Great Basin Naturalist 35: 378. 1976. Scrophulariaceae. Kane Co., sse Canaan Peak, Welsh & Welsh 12821, 1975 (BRY!;UT!;NY!;UTC!). Penstemon azureus var. ambiguus Gray Syn. Fl. N. Amer. 2: 272. 1886. Scrophulariaceae. = P. sepalulus A. Nels. Utah Co., near Provo, Watson 786, 1869 (US!;NY!). Penstemon bracteatus Keck Leafl. W. Bot. 1: 82. 1934. Scrophulariaceae. Garfield Co., Red Canyon, East- wood & Howell 783, 1933 (CAS!). Penstemon caespitosus Nutt. e.\ Gray ssp. perbrevis Pennell Contr. U.S. Natl. Herb. 20: 375. 1920. Scro- phulariaceae. Carbon Co., near Castle Gate, Pennell 6138, 1915 (NY!:US!). Penstemon caespitosus Nutt. ex Gray var. suffruti- cosus Gray Syn. Fl. N. Amer. 2: 270. 1878. Scrophula- riaceae. = P. tiisharensis N. Holmgren Beaver Co., near Beaver, Palmer sn, 1877 (ISC!). Penstemon carnosus Pennell in Graham Ann. Car- negie Mus. 25: 329. 1937. Scrophulariaceae. Emery Co., San Rafael Swell, Cottam 5229, 1935 (UT!;NY!;UTC!). Penstemon comarrhenus Gray Proc. Amer. Acad. 12: 81. 1876. Scrophulariaceae. Garfield (?) Co., Aquarius Plateau, Ward 462, 1875 (NY!;US!). June 1982 Welsh: Utah Plant Types 179 Penstemon coticinnus Keck Amer. Midi. Naturalist 23: 608. 1940. Scrophulariaceae. Millard Co., Tunnel Springs Range, Cottam 5634, 1933 (CAS!;US!;UT!;NY!). Penstemon confertus var. aberrans Jones Proc. Calif. Acad. II, 5: 715. 1895. Scrophulariaceae. = P. procerus var. aberrans (Jones) A. Nels. Wasatch Co., Soldier Sum- mit, Jones 5601i, 1894 (POM!;US!). Penstemon confusus Jones Zoe 4: 280. 1893. Scro- phulariaceae. Juab Co., Detroit, Jones sn, 1891 (POM!;CAS!;BRY!). Penstemon crandallii A. Nels. ssp. atratus Keck Bull. Torrey Bot. Club 64: 370. 1937. Scrophulariaceae. = F. crandallii var. atratus (Keck) N. Holmgren San Juan (?) Co., La Sal Mts., Jones sn, 1914 (CAS!;US!; POM!;NY!;BRY!). Penstemon cyananthus Hook. ssp. compactus Keck Amer. Midi. Naturalist 23: 615. 1940. Scrophulariaceae. = P. compactus (Keck) Crosswhite Cache Co., Mt. Naomi, Maguire 16148, 1938 (UTC!). Penstemon cyananthus Hook. ssp. longiflorus Pennell Contr. U.S. Natl. Herb. 20: 353. 1920. Scrophulariaceae. = P. longiflorus (Pennell) S. Clark Beaver Co., near Bea- ver, Palmer 376, 1877 (US!;NY!). Penstemon dolius Jones ex Pennell Contr. U.S. Natl. Herb. 20: 341. 1920. Scrophulariaceae. Tooele Co., Deep Creek Mts., Jones sn, 1891 (POM!;BRY!). Penstemon dolius Jones var. duchesnensis N. Hol- mgren Brittonia 31: 219. 1979. Scrophulariaceae. Duch- esne Co., e Duchesne, Holmgren et al, 8762, 1978 (NY!;BRY!;UTC!). Penstemon eatonii Gray Proc. Amer. Acad. 8: 395. 1872. Scrophulariaceae. Utah Co., Provo Canyon, Wat- son 776, 1869 (US!;NY!). Penstemon eatonii Gray var. undosus Jones Proc. Ca- lif. Acad. II, 5: 715. 1895. Scrophulariaceae. Syn: P. ea- tonii ssp. undosus (Jones) Keck Washington Co., St. George, Jones 51101h, 1894 (POM!;US!;NY!). Penstemon garrettii Pennell Contr. U.S. Natl. Herb. 20: .3.53. 1920. Scrophulariaceae. = P. scariosus Pennell sens. lat. Wasatch Co., near Midway, Carlton & Garrett 6697, 1905 (NY!;US!). Penstemon glaber var. utahensis Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 217. 1871. Scrophula- riaceae. = P. subglaber Rydb. Summit Co., Uinta Mts, Watson 771, 1869 (US!;NY!). Penstemon goodrichii N. Holmgren Brittonia 30: 416. 1978. Scrophulariaceae. Uintah Co., e Lapoint, Hol- mgren et al 8760, 1978 (NY!;BRY!;UTC!). Penstemon grahamii Keck in Graham Ann. Carnegie Mus. 26: 331. 1937. Scrophulariaceae. Uintah Co., Sand Wash, Graham 7883, 1933 (CM!). Penstemon heterophyllus var. latifolius Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 222. 1871. Scro- phulariaceae. = P. platijphyllus Rydb. Salt Lake Co., Cottonwood Canyon, Watson 787, 1869 (US!). Penstemon humilis Nutt. in Gray var. brevifolius Gray Syn. Fl. N. Amer. 2(1): 267. 1878. Scrophula- riaceae. Salt Lake Co., Cottonwood Canyon, Watson 781, 1869 (US!;NY!). Penstemon jonesii Pennell (hybrid) Contr. U.S. Natl. Herb. 20: 338. 1920. Scrophulariaceae. = P. eatonii x P. leiophyllus Washington Co., Springdale, Jones 5250, 1894 (US!). Penstemon laevis Pennell Contr. U.S. Natl. Herb. 20: .347. 1920. Scrophulariaceae. Washington Co., Spring- dale, Jones 5250, 1894 (US!;POM!). Penstemon leiophyllus Pennell Contr. U.S. Natl. Herb. 20: 346. 1920. Scrophulariaceae. Garfield Co., Mammoth Creek, Jones 6026b, 1894 (US!). Penstemon lentus Pennell ssp. albiflorus Keck Amer. Midi. Naturalist 23: 616. 1940. Scrophulariaceae. San Juan Co., 8 mi w Blanding, Porter 1801, 19.39 (RM!;NY!;UTC!). Penstemon leonardii Rydb. Bull. Torrey Bot. Club 40: 483. 1913. Scrophulariaceae. Wasatch Mts., Leonard sn, 1884 (NY!). Penstemon leptanthus Pennell Contr. U.S. Natl. Herb. 20: 339. 1920. Scrophulariaceae. Sanpete Co., Twelve Mile Creek, Ward 280, 1875 (US!). Penstemon linarioides Gray in Torr. var. sileri Gray Syn. Fl. N. Amer. 2(1): 270. 1878. Scrophulariaceae. Kane (?) Co., S. Utali, Siler sn, 1873 (NY!). Penstemon mucronatus N. Holmgren Brittonia 31: 234. 1979. Scrophulariaceae. Daggett Co., s. Manila, Holmgren et al. 8447, 1978 (NY!;BRY!;UTC!). Penstemon nanus Keck Amer. Midi. Naturalist 23: 607. 1940. Scrophulariaceae. Millard Co., Desert Experi- mental Range, Plummer 7313, 1939 (CAS!;BRY!; UT!;NY!;UTC!). Penstemon navajoa N. Holmgren Brittonia 30: 419. 1978. Scrophulariaceae. San Juan Co., Navajo Mt., N. & P Holmgren 8587, 1977 (NY!;BRY!;UTC!). Penstemon obtusifolius Pennell Contr. U.S. Natl. Herb. 20: 370. 1920. Scrophulariaceae. = P. humilis var. obtusifolius (Pennell) Reveal Washington Co., Spring- dale, Jones 5249am, 1894 (US!;POM!;NY!). Penstemon ophianthus Pennell Contr. U.S. Natl. Herb. 20: 343. 1920. Scrophulariaceae. Syn: P. jamesii ssp. ophianthus (Pennell) Keck Wayne Co., Thurber, Jones 5708, 1894 (US!;POM!;NY!). Penstemon palmeri Gray ssp. eglandulosus Keck Amer. Midi. Naturalist 18: 797. 19.37. Scrophulariaceae. = P. palmeri var. eglandulosus (Keck) N. Holmgren Kane Co., 2.5 mi n Kanab, Maguire et al. 12279, 1935 (RM!;NY!;UTC!). Penstemon parvus Pennell Contr. U.S. Natl. Herb. 20: 345. 1920. Scrophulariaceae. Garfield Co., Aquarius Plateau, Ward 546, 1875 (US!). Penstemon patricus N. Holmgren Brittonia 31: 238. 1979. Scrophulariaceae. Juab Co., Thorns Creek Canyon, Holmgren et al. 9018, 1978 (NY!;BRY!;UT!;UTC!). Penstemon phlogifolius Greene Leaf!. Bot. Obs. & Grit. 1: 164. 1906. Scrophulariaceae. = P. watsonii Gray Carbon Co., Castle Gate, Jones 5486s, 1894 (US!;POM!).' Penstemon platijphyllus Rydb. Bull. Torrey Bot. Club 36: 690. 1909. Scrophulariaceae. Syn: P. heterophyllus var. latifolius Wats. Salt Lake Co., Cottonwood Canyon, Watson 787, 1869 (US!). Penstemon pseudohumilis Jones Contr. W. Bot. 12: 65. 1908. Scrophulariaceae. = P. marcusii (Keck) N. Holmgren? Carbon Co., Price, Jones sn, 1898 (POM!). Penstemon pumilus var. thompsoniae Gray Syn. Fl. N. Amer. 2(1): 269. 1878. Scrophulariaceae. Syn: P. caespitosus var. thompsoniae (Gray) A. Nels. = P. thompsoniae (Gray) Rydb. Kane Co., Kanab, Thompson sn, 1872 (GH). 180 Great Basin Naturalist Vol. 42, No. 2 Penstemon scariosus Pennell Contr. U.S. Natl. Herb. 20: 353. 1920. Scrophulariaceae. Syn: P. garrettii Pennell Sanpete Co., Musinia Peak, Tidestrom 568, 1907 (US!). Penstemon tidestromii Pennell Contr. U.S. Natl. Herb. 20: 379. 1920. Scrophulariaceae. Sanpete Co., San Pitch Mts., Tidestrom 1296, 1908 (US!). Penstemon uintahensis Pennell Contr. U.S. Natl. Herb. 20: ,350. 1920. Scrophulariaceae. Uintah Co., Dyer Mine, Goodding 1221, 1902 (US!;RM!;BRY!;NY!). Penstemon utahensis Eastw. Zoe 4: 124. 1893. Scro- phulariaceae. San Juan Co., n Monticello, Eastwood sn, 1892 (CAS!;US!). Penstemon wardii Gray Proc. Amer. Acad. 12: 82. 1876. Scrophulariaceae. Sevier Co., near Glenwood, Ward 162, 1875 (US!;NY!). Petalonyx parryi Gray Proc. Amer. Acad. 10: 77. 1874. Loasaceae. Washington Co., St. George ("within a stones throw of the great Mormon temple"). Parry 75, 1874 (NY!;BRY!;ISC!). Petalostemon flavescens Wats. Amer. Naturalist 7: 299. 1873. Fabaceae. = Dalea flavescens (Wats.) Welsh Kane Co., "Kanab," Thompson sn, 1871 (US!). Peteria thompsonae Wats. Amer. Naturalist 7: 300. 1873. Fabaceae. Kane Co., Kanab, Thompson sn, 1872 (GH;US!;NY!;BRY!). Peucedanum bicolor Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 129. 1871. Apiaceae. = Lornatium bicolor (Wats.) Coult. & Rose Summit Co., Parleys Park, Watson 467, 1869 (US!;NY!). Peucedanum graveolens Wats. Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot. 5: 128. 1871. Apiaceae. = Lo- rnatium nuttallii (Gray) Macbr. Wasatch Mts., Watson 463, 1869 (US!;NY!). Peucedanum juniperinum Jones Contr. W. Bot. 8: 29. 1898. Apiaceae. = Lomatiiim juniperinum (Jones) Coult. & Rose Summit Co., Coalville, Jones sn, 1889 (POM!;US!;BRY!). Peucedanum lapidosum Jones Zoe 2: 246. 1891. Apiaceae. = Cymopterus longipes Wats. Summit Co., Echo, Jones sn, 1890 (US!;RM!;POM!). Peucedanum millefolium Wats. Rep. U.S. Explor. 40th Parallel, Bot. 5: 129. 1871. Apiaceae. = Lornatium grayi Coult. & Rose Davis Co., Antelope Island, Watson 466, 1869 (NY!;US!). Peucedanum parryi Wats. Proc. Amer. Acad. 11: 143. 1876. Apiaceae. = Lomatium parryi (Wats.) Macbr. Washington (?) Co., Southern Utah, Parrv 85, 1874 (NY!). Peucedanum triternatum var. platycarpum Torr. in Standb. Explor. Great Salt Lake 389. 1852. Apiaceae. = Lomatium triternatum var. platycarpum (Torr.) Cronq. Great Salt Lake, Stansbury sn, 18.50 (NY!). Phaca mollissima var. utahensis Torr. in Stansb. Ex- plor. Great Salt Lake 385. 1852. Fabaceae. = Astragalus utahensis (Torr.) T. & G. Tooele Co., w shore Stansbury Island, Great Salt Lake, Stansury sn, 1850 (NY!). Phaca pardalina Rydb. N. Amer. Fl. 24: 352. 1929. Fabaceae. = Astragalus pardalinus (Rydb.) Barneby Emery Co., Cedar Mt., Jones sn, 1915 (NY!;POM!). Phacelia argillacea Atwood Phytologia 26: 437. 1973. Hydrophyllaceae. Utah Co., Spanish Fork Canyon, At- wood et al. 3091, 1971 (BRY!;UT!;NY!;US!;UTC!). Phacelia cephalotes Gray Proc. Amer. Acad. 10: 325. 1875. Hydrophyllaceae. Washington Co., near St. George, Parry 179, 1874 (NY!;US!;BRY!;ISC!). Phacelia demissa Gray var. heterotricha J.T. Howell Amer. Midi. Naturalist 29(1): 8. 1943. Hydrophyllaceae. Piute Co., Marysvale, Jones 5388o, 1894 (POM!;US!). Phacelia foetida Goodding Bot. Gaz. 37: 58. 1904. Hydrophyllaceae. = P. palmeri Torr. Washington Co., Diamond Valley, Goodding 833, 1902 (RM!). Phacelia howelliana Atwood Rhodora 74: 456. 1972. Hydrophyllaceae. San Juan Co., Bluff, Atwood 2454, 1970 (BRY!;RM!;NY!;US!;UTC!). Phacelia incana Brand Beit. z. Kenntnis der Hydro- phyll. 8. 1911. Hydrophyllaceae. Tooele Co., Dugway, Jones sn, 1891 (POM!;RM!;NY!). Phacelia indecora J.T. Howell Amer. Midi. Naturalist 29: 12. 1943. Hydrophyllaceae. San Juan Co., Bluff, Jones sn, 1919 (CAS!). Phacelia mammillariensis Atwood Phytologia 26: 437. 1973. Hydrophyllaceae. Kane Co., e Glen Canyon City, Welsh & Atwood 9809, 1970 (BRY!). Phacelia nudicaulis Eastw. Zoe 4: 123. 1893. Hydro- phyllaceae. = P. demissa Gray var. demissa Grand Co., near Moab, Eastwood sn, 1892 (CAS!;POM!;US!). Phacelia orbicularis Rydb. Bull. Torrey Bot. Club 40: 479. 1913. Hydrophyllaceae. = P. corrtigata A. Nels. Wayne Co., Marvine Laccolite, Jones 5663, 1894 (US!;NY!). Phacelia palmeri Torr. in Wats. Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot. 5: 251. 1871. Hydrophyllaceae. Syn: P. foetida Goodding Washington Co., near St. George, Palmer 4, 1870 (NY!;US!). Phacelia pinetorum Jones Zoe 4: 279. 1893. Hydro- phyllaceae. = Eucrypta micrantha Torr. Tooele or Juab Co., Deep Creek Mts., Jones sn, 1891 (POM!;CAS!). Phacelia pulchella Gray Proc. Amer. Acad. 10: 326. 1875. Hydrophyllaceae. Washington Co., near St. George, Parry 182, 1874 (GH;POM!;NY!;US!;BRY!). Phacelia pulchella Gray var. sabulonum J.T.Howell Amer. Midi. Naturalist 29: 12. 1943. Hydrophyllaceae. = P. pulchella Gray Kane Co., Kaiparowits Plateau, Tompkins sn, 1939 (CAS!). Phacelia rafaelensis Atwood Rhodora 74: 454. 1972. Hydrophyllaceae. Wayne Co., Capitol Reef National Monument, Atwood & Higgins 1834, 1969 (BRY!; CAS!;RM!;NY!;US!;UTC!). Phacelia rotundifolia Torr. ex Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 253. 1871. Hydro- phyllaceae. Southern Utah, Palmer sn, 1870 (NY!;US!). Phacelia utahensis Voss Bull. Torrey Bot. Club 64: 135. 1937. Hydrophyllaceae. Sanpete Co., Gunnison, Jones sn, 1910 (POM!). Phlox austromontana Gov. Contr. U.S. Natl. Herb. 4: 151. 1893. Polemoniaceae. Washington Co., Beaverdam Mts., Bailey 1944, 1891 (US!). Phlox austromontana Gov. var. prostrata E. Nels. Rev. W. N. Amer. Phloxes 19. 1899. Polemoniaceae. Syn: P. jonesii Wherry Washington Co., Silver Reef, Jones 5163y-z, 1894 (NY!;US!;RM!;POM!;BRY!). Phlox caesia Eastw. Leafl. W. Bot. 2: 54. 1937. Po- lemoniaceae. = P. gladiformis (Jones) E. Nels. Garfield Co., Red Canyon, Eastwood & Howell 752, 1933 (CAS!). Phlox canescens T. & G. in Beckwith Rep. U.S. Ex- plor. & Surv. R.R. Pacific 2: 122. 1857. Polemoniaceae. = P. hoodii var. canescens (T. & G.) Peck Tooele Co., s Great Salt Lake, Beckwith 4, 1854 (NY!). June 1982 Welsh: Utah Plant Types 181 Phlox cluteana A. Nels. Amer. Botanist 28: 24. 1922. Polemoniaceae. San Juan Co., Navajo Mt., Clute 18, 1919(?). Phhx densa Brand Pflanzenr. 4. Fam. 250. 83. 1907. Polemoniaceae. = P. austromontana Gov. Beaver Co., Frisco, Jones 2021, 1880 (US!;POM!;NY!;UTC!). Phlox grahamii Wherry Brittonia 5: 63. 1943. Polemo- niaceae. = P. longifolia Nutt. Uintah Co., Sand Wash, Graham 7884, 1933 (CM!). Phlox jonesii Wherry Notul. Nat. Acad. Nat. Sci. Philadelphia 146: 8. 1944. Polemoniaceae. = P. austro- montana var. prostrata E. Nels. Washington Co., Zion Canyon, Jones sn, 1923 (US!;CAS!;POM!). Phlox longifolia Nutt. var. gladiformis Jones Proc. Calif. Acad. II, 5: 711. 1895. Polemoniaceae. = P. gladi- formis (Jones) E. Nels. Iron Co., Cedar Canyon, Jones 5208c, 1894 (POM!;US!). Physaria acutifolia Rydb. var. purpurea Welsh & Re- veal Great Basin Naturalist 37: 345. 1977. Brassicaceae. Grand Co., near Sego, Welsh 6902, 1968 (BRY!). Physaria chambersii Rollins var. membranacea Roll- ins Rhodora 41: 404. 1939. Brassicaceae. = F. lepidota var. membranacea (Rollins) Rollins Garfield Co., Red Canyon, Rollins & Chambers 2448, 1938 (GH; RM!;US!;NY!). Physaria grahamii Morton in Graham Ann. Carnegie Mus. 26: 220. 1937. Brassicaceae. Uintah Co., Chandler Canyon, Graham 9976, 1935 (US!;CM!). Physaria lepidota Rollins Brittonia 33: 335. 1981. Brassicaceae. Kane Co., between Kanab and Zion, Roll- ins & Rollins 79198, 1979 (GH). Physaria newberryi Gray var. racemosa Rollins Brit- tonia 33: 339. 1981. Brassicaceae. Washington Co., 13.5 mi s St. George, Holmgren et al. 9183, 1979 (GH). Pinus contort a Dougl. ex Loud. var. latifolia Engelm. ex Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 331. 1871. Pinaceae. = P. contorta Dougl. Summit Co., Uinta Mts., Watson 1110, 1869 (?). Plantago major L. var. pachyphylla Pilger Feddes Repert. 18: 277. 1922. Plant aginaceae. = P. major L. Salt Lake Co., Salt Lake City, Jones 1030, 1879 (POM!;UT!). Plantago myosuroides Rydb. Mem. New York Bot. Card. 1: 369. 1900. Plantaginaceae. = P. elongata Pursh Salt Lake Co., Salt Lake Valley, near mouth of Jordan River, Watson 749, 1869 (US!). Platystemon remotus Greene Pittonia 5: 190. 1903. Papaveraceae. = P. californicus Benth. Washington Co., near St. George, Parry 8, 1874 (GAS!;1SC!). Platystemon rigidulus Greene Pittonia 5: 167. 1903. Papaveraceae. = P. californicus Benth. Washington Co., near St. George, Parry 8a, 1874 (CAS!). Platystemon terminii Fedde Pflanzenr. 4. Fam. 104: 128. 1909. Papaveraceae. = F. californicus Benth. Washington Co., Diamond Valley, Jones 5121, 1894 (POM!;US!). Pleurophragma platypodum Rydb. Bull. Torrey Bot. Club 34: 4.34. 1907. Brassicaceae. = Thelypodium integ- rifolium var. gracilipes Robins. Grand Co., Moab, Jones sn, 1891 (NY!;CAS!;RM!;US!;BRY!). Poa eatonii Wats. Rep. U.S. Geol. Explor. 40th Paral- lel, Bot. 5: 386. 1871. Poaceae. = F. fendleriana (Steud.) Vasey Salt Lake Co., Cottonwood Canyon, Eaton 1891, 1869 (NY!). Poa festucoides Jones Proc. Calif. Acad. II, 5: 723. 1895. Poaceae. = Festuca thurberi Vasey Garfield Co., Mt. Ellen, Jones 5671, 1894 (POM!;US!). Poa longiligula Scribn. & Will. USDA Div. Agrostol. Circ. 9: 3. 1899. Poaceae. = P. fendleriana (Steud.) Vasey sens. lat. Washington Co., Silver Reef, Jones 5149at, 1894 (RM!;NY!;US!). Poa scabriuscula Williams USDA Div. Agrostol. Circ. 10: 4. 1899. Poaceae. = P. fendleriana (Steud.) Vasey Sevier Co., s Glenwood, Ward 136, 1895 (US!). Polemonium albiflorum Eastw. Bot. Gaz. 37: 437. 1904. Polemoniaceae. = P. foliosissimum var. alpinum Brand. Carbon Co., Scofield, Harkness sn, 1902 (CAS!;RM!). Polemonium foliosissimum Gray ssp. albiflorum var. alpinuum Brand Pflanzenr. 4. Fam. 250: 34. 1907. Po- lemoniaceae. Syn: P. albiflorum Eastw.; P. foliosissimum ssp. albiflorum (Eastw.) Brand Salt Lake Co., Alta, Jones 1114, 1879 (NY!;POM!;UT!). Polygala acanthoclada Gray Proc. Amer. Acad. 11: 73. 1876. Polygalaceae. Syn: P. acanthoclada var. in- tricata Eastw. San Juan Co., San Juan R., Brandegee sn, 1873 (NY!). Polygala acanthoclada Gray var. intricata Eastw. Proc. Calif. Acad. II, 6: 283. 1896. Polygalaceae. = P. acanthoclada Gray San Juan Co., Willow Creek, East- wood 10, 1895 (CAS!). Polygonum minimum Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 315. 1871. Polygonaceae. Summit Co., Uinta Mts., Watson 1058, 1869 (US!). Polygonum utahense Brenckle & Cottam Bull. Univ. Utah 30: 3. 1940. Polygonaceae. Garfield Co., 6 m n Es- calante, Cottam 6507, 1935 (UT!;US!;BRY!;CAS!; RM!;UTC!). Potamogeton pusillus L. var. vulgaris subvar. inter- ruptus J.W. Robbins Rep. U.S. Geol. Explor. 40th Paral- lel, Bot. 5: 338. 1871. Potamogetonaceae. = P. pusillus var. pusillus Summit Co., Parleys Park, Silver Cr., Wat- son 1137, 1869 (NY!). Potentilla dissecta var. decurrens Wats. Proc. Amer. Acad. 8: 557. 1873. Rosaceae. Syn: P. decurrens (Wats.) Rydb. = P. ovina var. decurrens (Wats.) Welsh & John- ston Summit Co.?, Uinta Mts., Watson 329, 1869 (US!). Potentilla diversifolia var. pinnatisecta Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 87. 1871. Ro- saceae. = P. ovina Macoun Summit Co., Uinta Mts., Watson 331, 1869 (NY!). Potentilla modesta Rydb. N. Amer. Fl. 22: 331. 1908. Rosaceae. = P. concinna var. modesta (Rydb.) Welsh & Johnston Piute Co., Mt. Barrette, near Marysvale, Ryd- berg & Carlton 7261, 1905 (NY!). Potentilla paucijuga Rydb. N. Amer. Fl. 22: 348. 1908. Rosaceae. = P. pensylvanica var.- paucijuga (Rydb.) Welsh & Johnston Grand (?) Co., La Sal Mts., Purpus 251, 1899 (NY!). Potentilla pectinisecta Rydb. Bull. Torrey Bot. Club 24: 7. 1897. Rosaceae. = P. gracilis Dougl. Salt Lake Co., Salt Lake City, Jones 1765, 1880 (RM!;NY!;BRY!). Potentilla proxima Rydb. N. Amer. Fl. 22(4): 339. 1908. Rosaceae. = F. concinna var. proxima (Rydb.) Welsh & Johnston Piute Co, s Belknap Peak, Rydberg & Carlton 7369, 1905 (NY!). 182 Great Basin Naturalist Vol. 42, No. 2 Potentilla sabulosa Jones Proc. Calif. Acad. Sci. II, 5: 680. 1895. Rosaceae. = Ivesia sabulosa (Jones) Keck Garfield Co., Sevier River, Jones 6032, 1894 (US!; POM!;CAS!;RM!;NY!). Potentilla wardii Greene Leafl. Bot. Obs. & Crit. 2: 138. 1911. Rosaceae. = P. hippiana Lehm. Wayne Co., Thousand Lake Mt., Ward sn, 1875 (US!). Primula incana Jones Proc. Calif. Acad. II, 5: 705. 1895. Primulaceae. Garfield Co., Sevier River, Jones 5312av, 1894 (POM!). Primula maguirei L.O. Williams Amer. Midi. Natu- ralist 7: 747. 1936. Primulaceae. Cache Co., Logan Can- yon, Maguire & Maguire 3650, 1932 (M0;UT!;UTC!). Primula specuicola Rydb. Bull. Torrey Bot. Club 40: 461. 1913. Primulaceae. San Juan Co., near Bluff, Ryd- berg9882, 1911 (NY!;US!;UT!). Prosartes trachycarpa Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 344. 1871. Liliaceae. = Disponim trachycarpum Wats. Summit Co., Parleys Park, Watson 1166,1869 (NY!). Psathyrotes pilifera Gray Proc. Amer. Acad. 19: 50. 1883. Asteraceae. Kane Co., Kanab, Thompson sn, 1872 (GH). Pseudocymoptems tidestromii Coult. & Rose Contr. U.S. Natl. Herb. 12: 447. 1909. Apiaceae. = Cymopterus lemmonii (Coult. & Rose) Dorn Sevier Co., Mt. Terrell, Tidestrom 1811, 1908 (US!). Pseudocymoptems versicolor Rydb. Fl. Rocky Mts. 623, 1064. 1917. Apiaceae. = Cymopterus lemmonii (Coult. & Rose) Dorn Garfield Co., Aquarius Plateau, Rydberg & Carlton 7426, 1905 (NY!). Pseudopteryxia longiloba Rydb. Bull. Torrey Bot. Club 40: 72. 1913. Apiaceae. = Cymopterus hendersonii (Coult. & Rose) Cronq. San Juan Co., Abajo Mts., Ryd- berg & Garrett 9761, 1911 (NY!;UT!). Pseudotsuga globosa Flous Bull. Soc. Hist. Nat. Tou- louse 66: 334. 1934. Pinaceae. = P. menziesii (Mirb.) Franco Salt Lake Co., Big Cottonwood Canyon, Ryd- berg & Carlton 6587, 1905 (?). Psoralea castorea Wats. Proc. Amer. Acad. 14: 291. 1879. Fabaceae. Beaver Co. (more likely Beaverdam, Arizona), near Beaver City, Palmer 96, 1877 (US!;NY!;BRY!;ISC!). Psoralea epipsila Bameby Leafl. W. Bot. 3: 193. 1943. Fabaceae. Kane Co., 17 mi e Kanab, Ripley & Barneby 4832, 1942 (CAS!). Psoralea juncea Eastw. Proc. Calif. Acad. 11, 6: 286. 1897. Fabaceae. San Juan Co., Epsom Creek, Eastwood 21, 1895 (CAS!;US!;NY!). Psoralea mephitica Wats. Proc. Amer. Acad. 14: 291. 1879. Fabaceae. Washington Co., near St. George, Pal- mer 97, 1877 (US!;NY!;BRY!;1SC!). Psoralea pariensis Welsh & Atwood in Welsh, At- wood, & Reveal Great Basin Naturalist 35: 353. 1975. Fabaceae. Garfield Co., Bryce Canyon, Welsh & Mur- dock 12859, 1975 (BRY!;UT!;NY!;US!;UTC!;ISC!). Psoralea rdfaelensis Jones Contr. W. Bot. 18: 41. 1933. Fabaceae. = P. aromatica Payson Grand Co., LaS- al Mts., Jones sn, 1914 (US!;CAS!). Psoralea rafaelensis Jones var. magna Jones Contr. W. Bot. 18: 41. 1933. Fabaceae. = P. aromatica Payson Emery Co., San Rafael Swell, Jones sn, 1914 (CAS!). Psoralea stenophylla Rydb. Bull. Torrey Bot. Club 40: 46. 1913. Fabaceae. = F. lanceolata var. stenophylla (Rydb.) Toft & Welsh Grand Co., Wilson Mesa, Rydberg & Garrett 8367, 1911 (NY!). Psoralea stenostachys Rydb. Bull. Torrey Bot. Club 40: 46. 1913. Fabaceae. = P. lanceolata var. stenos- tachys (Rydb.) Welsh Tooele Co., Government Well, Jones 6221, 1900 (NY!;US!;CAS!;POM!;RM!;BRY!). Ptelea neglecta Greene Contr. U.S. Natl. Herb. 10: 71. 1906. Rutaceae. = P. trifoliata ssp. pallida var. lutes- cens (Greene) V.K. Bailey Kane Co., near Kanab, Weth- erill sn, 1897 (CAS!). Pterchiton occidentale Torr. & Frem. in Frem. Rep. Exped. Rocky Mts. 318. 1845. Chenopodiaceae. = Atri- plex canescens (Pursh) Nutt. Davis (?) Co., Great Salt Lake, Fremont sn, 1843 (NY!). Ptilocalais macrolepis Rydb. Bull. Torrey Bot. Club 38: 11. 1911. Asteraceae. = Microseris nutans (Geyer) Schulz-bip Salt Lake Co., Salt Lake City, Garrett 182, 1903 (US!;NY!;BRY!). Pyrrocoma cheiranthifolia Greene Leafl. Bot. Obs. & Crit. 2: 47. 1910. Asteraceae. = Haplopappus dementis (Rydb.) Blake Sanpete Co., w. Ephriam, Tidestrom 534, 1907 (US!). Pyrrocoma lapathifolia Greene Leafl. Bot. Obs. & Crit. 2: 13. 1909. Asteraceae. = Haplopappus integri- folius Gray? Utah, Ward 596, 1875 (US!). Pyrrocoma subcaesia Greene Leafl. Bot. Obs. & Crit. 2: 12. 1909. Asteraceae. = Haplopappus dementis (Rydb.) Blake Aarfield Co., Panguitch Lake, Jones 6005, 1894 (US!;RM!;POM!;NY!;BRY!). Quercus eastwoodiae Rydb. Bull. New York Bot. Card. 2: 210. 1901. Fagaceae. = Q. gambelii Nutt. San Juan Co., Butler Wash, Eastwood 141, 1895 (NY!;US!). Quercus stellata var. utahensis A. DC. Prodr. 16(2): 22. 1864. . = Q. gambdii Nutt. Utah? (between Salt Lake and Sierras) Beckwith (?) sn, 1854?(G). Ranunculus acriformis Gray var. aestivalis L. Benson Amer. Midi. Naturalist 40: 43. 1948. Ranunculaceae. Garfield Co., 8.3 mi n Panguitch, Benson 13421, 1948 (POM;RM;BRY!;UT!;UTC!). Ranunculus alismaefolius Geyer ex Benth. var. mon- tanus Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 7. 1871. Ranunculaceae. Summit (?) Co., Uinta Mts., Watson 18, 1969 (US!). Ranunculus andersonii Gray var. tenellus Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 7. 1871. Ra- nunculaceae. = R. juniperinus Jones Salt Lake (?) Co., Pilot Rock Point, Salt Lake, Watson 17, 1869 (?). Ranunculus juniperinus Jones Proc. Calif. Acad. II, 5: 616. 1895. Ranunculaceae. Syn: R. andersonii var. te- nellus Wats. Washington Co., 18 mi w St. George, Jones 5011, 1894 (US!). Ranunculus multifidus Pursh var. repens Hook, in Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 8. 1871. . = R. gmdinii var. hookeri (D. Don) L. Benson Summit Co., Weber Valley, Watson 24, 1869 (US!). Ranunculus orthrorhynchus Hook. var. alpinus Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 9. 1871. Ranunculaceae. = R. adoneus var. alpinus (Wats.) L. Benson Salt Lake (?) Co., Wasatch Mts., Watson 30, 1869 (US!). Ranunculus orthrorhynchus Hook. var. platyphyllus Gray Proc. Amer. Acad. 21: 377. 1886. Ranunculaceae. Salt Lake (?) Co., Wasatch Mts., Watson 28, 1869 (US!). June 1982 Welsh: Utah Plant Types 183 Ranuticulus utahensis Rydb. Bull. Torrey Bot. Club 29: 158. 1902. Ranunculaceae. = R. inamoenus var. al- peaphiltis (A. Nels.) L. Benson Salt Lake Co., Alta, Jones 1130, 1879 (NY!;POM!;UTC!). Rhus nitens Greene Proc. Washington Acad. Sci. 8: 190. 1906. Anacardiaceae. = R. glabra L. Utah Co., Pro- vo, Jones 5612, 1894 (US!;RM!;NY!). Rhus utahensis Goodding Bot. Gaz. 37: 57. 1904. .\nacardiaceae. = R. trilobata var. simplicifolia (Greene) Barklev Washington Co., Diamond Valley, Goodding 832, 1902 (US!;RM!). Rhysopterus jonesii Coult. & Rose Contr. U.S. Natl. Herb. 7: 186. 1900. Apiaceae. = Cymopterus coulteri (Jones) Mathias Juab Co., Juab, Jones 1691, 1880 (US!;POM!;BRY!). Rites lacustre (Pers.) Poir. in Lam. var. lentutn Jones Proc. Calif. Acad. II, 5: 681. 1895. Saxifragaceae. = R. montigenum McClatchie Garfield Co., Henry Mts., Jones 56950, 1894 (POM!). Riddellia tagetina var. sparsiflora Gray Syn. Fl. N. Amer. 1(2): 318. 1884. Asteraceae. = Psilostrophe spar- siflora (Gray) A. Nels. Kane Co., (Southern Utah), Bish- op, Mrs. Thompson sn, 1872 (GH?). Rorippa integra Rydb. Bull. Torrey Bot. Club 29: 236. 1902. Brassicaceae. = R. curvipes var. integra (Rydb.) Stuckey Wasatch Mts., Watson 64, 1869 (US!;NY!). Rosa chrysocarpa Rydb. Bull. Torrey Bot. Club 44: 74. 1917. Rosaceae. = R. woodsii Lindl. San Juan Co., Allen Canyon, sw of Abajo Mts., Rydberg & Garrett 9312, 1913 (NY!). Rosa puberulenta Rydb. Fl. Rocky Mts. 443. 1917. Rosaceae. = R. woodsii Lindl. San Juan Co., Mon- tezimia Canyon, e of Monticello, Rydberg & Garrett 9705, 1913 (NY!). Rumex maritimus L. var. athrix St.John Rhodora 17: 79. 1915. Polvgonaceae. = R. maritimus L. Sevier Co., Vemiillion, Jones 58.39, 1894 (MO;GH;POM!). Rumex subalpina Jones Proc. Calif. Acad. II, 5: 720. 1895. Polvgonaceae. = R. pycnanthus Rech. f. Piute Co., Brigham Peak, Jones 5957, 1894 (POM!; US!;RM!;BRY!;NY!). Rumex utahensis Rech. f. Repert. Sp. Nov. 40: 298. 19.36. Polvgonaceae. = R. salicifolius Wein. Carbon Co., Kyime, Jones 5603, 1894 (US!). Salicornia utahensis Tidestr. Proc. Biol. Soc. Wash- ington 26: 13. 1913. Chenopodiaceae. = S. pacifica var. utahensis (Tidestr.) Munz Utah (?). Salix lutea Nutt. var. platyphylla Ball Bot. Gaz. 71: 4.30. 1921. Salicaceae. = S. rigida Muhl. Salt Lake Co., City Creek Canyon, Ball 1.336, 1908 (US!). Saxifraga caespitosa L. ssp. exaratoides var. purpusii Engler & Irmsch. Pflanzenr. 4. Fam. 117(1): 377. 1916. Saxifragaceae. = S. caespitosa var. minima Blankinship Grand (?) Co., LaSal Mts., Purpus 6642, 1899 (RM!). Schmaltzia affinis Greene Leafl. Bot. Obs. & Crit. 1: 135. 1905. .\nacardiaceae. = Rhus trilobala var. sim- plicifolia (Greene) Barkeley Kane Co., Kanab, Jones 5286e, 1894 (POM!;US!). Schoencrambe pinnata Greene Pittonia 3: 127. 1896. Brassicaceae. = S. linifolia (Nutt.) Greene Utah, Ward 1875 (US!). Sclerocactus contorus Heil Cact. Succ. J. (U.S.) 51: 25. 1979. Cactaceae. San Juan Co., Canyonlands National Park, Heil sn, (UNM). Sclerocactus parviflorus Clover & Jotter Bull. Torrey Bot. Club 68: 419. 1941. Cactaceae. Syn: S. whipplei au- thors, not (Engelm. & Bigel) Britt. San Juan Co., Glen Canyon, Clover & Jotter 2398, 1940 (US!). Sclerocactus terrae-canyonae Heil Cact. Succ. J. (U.S.) 51: 26. 1979. Cactaceae. San Juan Co., without lo- cality, Heil sn, (UNM). Sclerocactus wrightiae L. Benson Cact. & Succ. J. (U.S.) 38: 55. 1966. Cactaceae. Emery Co., San Rafael Ridge, Benson & Benson 16595 (POM). Scrophularia utahensis Gandg. Bull. Soc. Bot. France 66: 219. 1919. Scrophulariaceae. = S. lanceolata Pursh? Cache Co., Linford sn, 1897 (?). Sedum meehanii Gray Proc. Amer. Acad. 16: 105. 1880. Crassulaceae. = ? S. lanceolatum Torr. Salt Lake Co., City Creek Canyon, Reading (GH). Selaginella utahensis Flowers Amer. Fern J. 39: 83. 1949. Selaginellaceae. Washington Co., s St. George, Cottam 5644, 1931 (UT!;BRY!). Selaginella watsonii Underwood Bull. Torrey Bot. Club 25: 127. 1898. Selaginellaceae. Salt Lake Co., Cot- tonwood Canyon, Watson 2370, 1869 (US!). Senecio aquariensis Greenm. Ann. Missouri Bot. Card. 3: 144. 1916. Asteraceae. = S. streptanthifolius Greene Garfield Co., Aquarius Plateau, Ward 505, 1875 (GH;US!). Senecio convallium Greenm. Ann. Missouri Bot. Card. 1: 266. 1914. Asteraceae. = S. canus Hooker Wavne Co., Rabbit Valley, Ward 704, 1875 (US!). Senecio dimorphophyllus Greene var. intermedius T.M. Barkley Trans. Kansas Acad. 65: 362. 1963. Aste- raceae. San Juan Co., LaSal Mts., Payson & Payson 4097, 1924 (MO;GH;RM!;UC). Senecio incurvus A. Nels. Univ. Wyoming Publ. Bot. 1: 141. 1926. Asteraceae. = S. spartioides T. & G. Wash- ington Co., Zion National Park, Nelson 9988, 1922 (RM!). Senecio jonesii Rydb. Bull. Torrey Bot. Club 27: 179. 1900. Asteraceae. = S. streptanthifolius Greene Salt Lake Co., Alta, Jones 1125, 1879 (NY!;POM!;UTC!). Senecio kingii Rydb. Bull. Torrey Bot. Club 37: 468. 1910. Asteraceae. = S. eremophilus var. kingii (Rydb.) Greenm. Salt Lake Co., Cottowood Canyon, Watson 676 (NY!;US!). Senecio lapidum Greenm. Ann. Missouri Bot. Gard. 4: 18. 1917. Asteraceae. = S. multilobatus T. & G. ex Gray Washington Co., Silver Reef, Jones 5163v, 1894 (NY!;BRY!). Senecio leonardii Rydb. Bull. Torrey Bot. Club 37: 468. 1910. Asteraceae. = S. streptanthifolius Greene Utah Co., American Fork Canyon, Leonard 143, 1884 (NY!). Senecio lugens Richards var. hookeri D.C. Eaton in Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 188. 1871. Asteraceae. = S. sphaerocephalus Greene Summit Co., Salt Lake City to Uintas, Watson 661, 1869 (US!;NY!). Senecio malmstenii Blake in Tidestr. Proc. Biol. Soc. Washington 36: 183. 1923. Asteraceae. = S. streptanthi- folius Greene Kane Co., Little Podunk Creek, Malmsten 131, 1916 (US!). Senecio multilobatus T. & G. ex Gray Mem. Amer. Acad. II, 4: 109. 1849. Asteraceae. Uintah Co., Uinta River, Fremont 549, 1845 (NY!). 184 Great Basin Naturalist Vol. 42, No. 2 Senecio pammelii Greenm. Ann. Missouri Bot. Card. 3: 118. 1916. Asteraceae. = S. streptanthif alius Greene Morgan Co., Peterson, Pammel & Blackwood .3870, 1902 (MO;GH). Senecio platylohus Rydb. Bull. Torrey Bot. Club 27: 181. 1900. Asteraceae. = S. streptanthifolius Greene Wasatch Mountains, Watson 671, 1869 (NY!). Senecio rubricaulis var. aphanactis Greenm. Ann. Missouri Bot. Card. 3: 174. 1916. Asteraceae. = S. strep- tanthifolius Greene Cache Co., Logan, Smith 2208, 1910 (UTC!). Senecio wardii Greene Pittonia 4: 116. 1900. Aste- raceae. = S. streptanthifolius Greene Sevier Co., Fish Lake Mt., Ward 332, 1875 (GH;US!). Shepherdia rotundifolia Parry Amer. Naturalist 9: 350. 1875. Elaeagnaceae. Kane Co., Valley of the Virgin, Siler sn, 1875 (US!;NDG!). SidaJcea crenulata A. Nels. Proc. Biol. Soc. Washing- ton 17: 93. 1904. Malvaceae. = S. neomexicana var. cre- nulata (A. Nels.) C.L. Hitchc. Juab Co., Juab, Goodding 1091, 1902 (RM!;BRY!;NY!;ISC!). Sieversia scapoidea A. Nels. in Coult. & Nels. New Man. Bot. Rocky Mts. 263. 1909. Rosaceae. = Geum rossii (R. Br.) Ser. Piute Co., w of Marysvale, Jones 5871, 1894 (RM!;POM!). Silene petersonii Maguire var minor Hitchc. & Ma- guire Univ. Washington Publ. Biol. 13: 38. 1947. Caryo- phyllaceae. Garfield Co., Red Canyon, Maguire 19550, 1940 (UTC!;NY!). Silene petersonii Maguire var. petersonii Madrono 6: 24. 1941. Caryophyllaceae. Sanpete Co., Wasatch Plateau, Maguire 20000, 1940 (UTC!;NY!;BRY!). Sisyrinchium radicatum Bickn. Bull. Torrey Bot. Club. 28: 576. 1901. Iridaceae. = S. demissiim Greene Washington Co., St. George, Palmer 456, 1877 (NY!;ISC!). Sitanion insulare J.G. Sm. USDA Div. Agrostol. Bull. 18: 14. 1899. Poaceae. = Agrositanion saxicola (Scribn. & Sm.)Bowden Tooele Co., Carrington Island, Watson 1338, 1869 (US!). Solidago garrettii Rydb. Bull. Torrey Bot. Club 37: 134. 1910. Asteraceae. = S. sparsi flora Gray Salt Lake Co., Big Cottonwood Canyon, Garrett 2041, 1906 (NY!;UT!). Solidago radulina Rydb. Bull. Torrey Bot. Club 31: 650. 1904. Asteraceae. = S. nana Nutt. Salt Lake Co., Cottonwood Canyon, Watson .558, 1869 (NY). Sonchus asper (L.) All. var. glanduliferus Garrett Spr. Fl. Wasatch ed. 3. 119. 1917. Asteraceae. = S. asper (L.) Hill Utah (?). Sophia leptostylis Rydb. Bull. Torrey Bot. Club .39: 325. 1912. Brassicaceae. = Descurainia californica (Gray) Schulz Salt Lake Co., Big Cottonwood Canyon, Rydberg & Carlton 6629, 1905 (NY!;US!;RM!). Sphaeralcea caespitosa Jones Contr. W. Bot. 12: 4. 1908. Malvaceae. Beaver (?) Co., Wah Wah, Jones sn, 1906 (POM!;CAS!;US!;BRY!;NY!). Sphaeralcea grossulariifolia (H. & A.) Rydb. var. moorei Welsh Great Basin Naturalist 40: 35. 1980. Mal- vaceae. Kane Co., Lake Powell, Welsh & Atwood 11597, 1972 (BRY!). Sphaeralcea leptophylla (Gray) Rydb. var. janeae Welsh Great Basin Naturalist 40: 36. 1980. Malvaceae. San Juan Co., White Rim road, Welsh 7085, 1968 (BRY!). Sphaeralcea psoraloides Welsh Great Basin Naturalist 40: 36. 1980. Malvaceae. Wayne Co., Salt Wash, Welsh 13348, 1977 (BRY!). Sphaeralcea subrhomhoidea Rydb. Bull. Torrey Bot. Club 40: .59. 1913. Malvaceae. = S. munroana (Dougl.) Spach Wasatch Co., Midway, Carlton & Garrett 6691, 1905 (NY!;US!). Sphaeromeria ruthiae Holmgren, Schultz, & Lowrey Brittonia 28: 257. 1976. Asteraceae. Washington Co., Zion Canyon, Holmgren et al. 16603, 1974 (UTC!; BRY!;UT!;US!;NY!). Sphaerostigma utahense Small Bull. Torrey Bot. Club 23: 191. 1896. Onagraceae. = Camissonia boothii ssp. alyssoides (H. & A.) Raven Beaver Co., Milford, Jones 1773, 1880 (NY!;POM!). Spiraea caespitosa Nutt. in T. & G. var. elatior Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 81. 1871. Rosaceae. = Petrophytum caespitosum (Nutt.) Rydb.. Box Elder Co., Raft R. Mts., Watson .307, 1869 (NY!). Stachys asperrima Rydb. Bull. Torrey Bot. Club 36: 682. 1909. Lamiaceae. = S. palustris L. Salt Lake Co., Jordan, Leonard 138, 1884 (NY!). Stanleya canescens Rydb. Bull. Torrey Bot. Club 29(1): 232. 1902. Brassicaceae. = S. pinnata var. pinnata Beaver Co., Frisco, Jones 1809, 1880 (NY!;POM!). Stenotus falcatus Rydb. Bull. Torrey Bot. Club 27: 616. 1900. Asteraceae. = Haplopappus acaulis var. gla- bratus (D.C. Eaton) Hall Washington Co. (s. Utah), Red Creek, Palmer 202, 1877 (US!;NY!;ISC!). Stenotus latifolius A. Nels. Bot. Gaz. 37: 266. 1904. Asteraceae. = Haplopappus acaulis var. glabratus (D.C. Eaton) H.M. Hall Utah Co., near Provo, Goodding 1111, 1902 (US!;NY!;BRY!;1SC!). Stipa arida Jones Proc. Calif. Acad. 11, 5: 725. 1895. Poaceae. Syn: S. mormonum Mez Piute Co., near Mary- svale, Jones 5377, 1894 (POM!;RM!;US!;NY!). Stipa mormonum Mez. Feddes Repert. 17: 209. 1921. Poaceae. = S. arida Jones Beaver Co., Milford, Jones 2106, 1880(US!frag;UT!;UTC!). Stipa parishii var. depauperata Jones Contr. W. Bot. 14: 11. 1912. Poaceae. = S. coronata var. depauperata (Jones) A.S. Hitchc. Juab Co., Detroit, Jones, 1891 (US!). Stipa pinetoTum Jones Proc. Calif. Acad. II, 5: 724. 1895. Poaceae. Garfield Co., Panguitch Lake, Jones 6023, 1894 (POM!;US!). Streptanthus crassicaulis Torr. in Stansbury Expl. Great Salt Lake, Append. D. Botany 383. 1852. Brassi- caceae. = Caulanthus crassicaulis (Torr.) Wats. Tooele Co., w shore of Great Salt Lake, Stansbury sn, 1850 (NY!). Suaeda intermedia Wats. Proc. Amer. Acad. 14: 296. 1879. Chenopodiaceae. Sevier Co., Glenwood, Ward 718, 1875 (US!). Swertia fritillaria Rydb. Bull. Torrey Bot. Club 40: 465. 1913. Gentianaceae. = S. perennis L. Salt Lake Co, Big Cottonwood Canyon, Garrett 1566, 1905 (NY!;UT!). Synthris laciniata (Gray) Rydb. ssp. ibahpensis Pen- nell Proc. Acad. Nat. Sci. Philadelphia 85: 92. 1933. Scrophulariaceae. = S. laciniata (Gray) Rydb. Juab Co., Mt. Ibapah, Stanton 1000, 1932 (PH). Synthris pinnatifida Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 227. 1871. Scrophulariaceae. Utah Co., American Fork Canyon, Watson 802, 1869 (US!;NY!). June 1982 Welsh: Utah Pla>jt Types 185 Synthris pinnatifida Wats. var. laciniata Gray Syn. Fl. N. Amer. 2(1): 286. 1878. Scrophulariaceae. = S. laciniata (Gray) Sevier Co., Fish Lake Mts., Ward 327, 1875 (US!). Tanacetum diversifolium D.C. Eaton Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 180. 1871. Asteraceae. = Sphaeromeria diversifolia Rydb. Utah Co., American Fork Canyon, Watson 632, 1869 (US!). Tetradymia linearis Rydb. Bull. Torrey Bot. Club 32: 130. 1905. Asteraceae. = T. canescens DC. Iron Co., Rock Creek, Palmer 264, 1877 (NY!). Tetradymia spinosa H. & A. var. longispina Jones Proc. Calif. Acad. II, 5: 698. 1895. Asteraceae. = T. ax- illaris var. longispina Jones Washington Co., St. George, Jones 5110, 1894 (POM!;US!;NY!;BRY!). Tetraneuris epunctata A. Nels. Bot. Gaz. 37: 275. 1904. Asteraceae. = Hymenoxys acaulis var. caespitosus (A. Nels.) Parker Uintah Co., Dyer Mine, Goodding 1236, 1902 (RM!;US!;UT!;NY!;BRY!;ISC!). Thalictrum duriusculum Greene Leafl. Bot. Obs. & Crit. 2: 92. 1910. Ranunculaceae. = T. alpinum L. Se- vier Co., near Fish Lake, Jones 5826a, 1894 (US!;POM!). Thelesperma subnudum Gray Proc. Amer. Acad. 10: 72. 1874. Asteraceae. Washington Co., near St. George, Parry 109, 1874 (US INY!;ISC!;BRY!). Thelypodiopsis argillacea Welsh & Atwood Great Ba- sin Naturalist 37: 95. 1977 Brassicaceae. Uintah Co., Big Pack Mountain, Atwood 6627, 1976 (BRY!). Thelypodiopsis bamebyi Welsh & Atwood in Welsh Brittonia 33: 300. 1981. Brassicaceae. Emery Co., San Rafael Swell, Welsh 20345, 1981 (BRY!;NY!;GH!;UTC!). Thelypodiutn elegans Jones Zoe 4: 265. 1893. Brassi- caceae. = Tlielypodiopsis elegans (Jones) Rydb. Grand Co., Westwater, Jones sn, 1891 (POM!;RM!;BRY!;NY!). Thelypodium macropetalum Rydb. Bull. Torrey Bot. Club 29: 233. 1902. Brassicaceae. = T. sagittatiim var. sagittatum Davis Co., Farmington, Jones 1841, 1881 (NY!;US!;CAS!;POM!;BRY!;UTC!). Thelypodium ovalifolium Rydb. Bull. Torrey Bot. Club 30: 253. 1903. Brassicaceae. = T. sagittatum var. ovalifolium (Rydb.) Welsh & Reveal Garfield Co., Pan- guitch Lake, Jones 6015e, 1894 (US!;POM!;NY!). Thelypodium palmeri Rydb. Bull. Torrey Bot. Club 34: 433. 1907. Brassicaceae. = T. sagittatum var. ovalifo- lium (Rydb.) Welsh & Reveal Washington Co., (s. Utah), Palmer 25, 1877 (NY!;US!;BRY!;ISC!). Thelypodium rollinsii Al-Shehbaz Contr. Gray Herb. II, 204: 97. 1973. Brassicaceae. Juab Co., 12 mi n Scipio, I. & N. Al- Shehbaz 6913, 1969 (GH). Thelypodium sagittatum (Nutt.) Endl. var. vermicu- laris Welsh & Reveal Great Basin Naturalist 37: 358. 1978. Brassicaceae. Sevier Co., 4 mi se Sigurd, Welsh & Atwood 11718, 1972 (BRY!). Thelypodium suffrutescens Rollins in Graham Ann. Carnegie Mus. 26: 224. 1937. Brassicaceae. = Glauco- carpum suffrutescens (Rollins) Rollins Uintah Co., Big Pack Mtn, Graham 8959, 1935 (GH;CM!;NY!). Thelypodium utahense Rydb. Bull. Torrey Bot. Club 29: 233. 1902. Brassicaceae. = Caulanthus lasiophyllus var. utahensis (Rydb.) Jeps. Washington Co., St. George, Jones 1648, 188(T(US!;UT!;POM!;NY!). Thelypodium wrightii Gray var. tenellum Jones Proc. Calif. Acad. II, 5: 622. 1895. Brassicaceae. = T. laxiflo- rum Al-Shehbaz Utah Co., Slate Canyon, Jones 5559, 1894 (POM!;US!;CAS!;RM!;NY!). Thlaspi fendleri Gray var. tenuipes Maguire Amer. Midi. Naturalist 24: 469. 1942. Brassicaceae. = T. mon- tanum var. montanum Sanpete Co., Mayfield Canyon, Maguire 19998, 1940 (UTC!). Thlaspi prolixum A. Nels. Amer. J. Bot. 32: 287. 1945. Brassicaceae. = T. montanum var. montanum Piute Co., Marysvale, Jones 5374, 1894 (RM!;POM!). Thysanocarpus trichocarpus Rydb. Bull. Torrey Bot. Club 30: 253. 1903. Brassicaceae. = T. curvipes Hook. Washington Co., Silver Reef, Jones 5163b, 1894 (US!). Tithonia argophylla D.C. Eaton Rep. U.S. Geol. Ex- plor. 40th Parallel, Bot. 5: 423. 1871. Asteraceae. = En- celiopsis argophylla (D.C. Eaton) A. Nels. Washington Co., St. George, Palmer sn, 1877 (US!). Townsendia annua Beaman Contr. Gray Herb. II, 183. 132. 1957. Asteraceae. San Juan Co., 1.5 mi n. Bluff, Maguire 13509, 1936 (UTC!). Townsendia aprica Welsh & Reveal Brittonia 20: 375. 1968. Asteraceae. Sevier Co., s Fremont Jet., Reveal & Welsh 721, 1966 (BRY!;CAS!). Townsendia dejecta A. Nels. Bot. Gaz. 37: 267. 1904. Asteraceae. = T. montana var. montana Uintah Co., Dyer Mine, Goodding 1283, 1902 (US!;RM!). Townsendia florifer (Hook.) Gray var. communis Jones Proc. Calif. Acad. II, 5: 697. 1895. Asteraceae. = r. florifer (Hook.) Gray Piute Co., Kingston, Jones 5322f, 1894 (POM!;US!;BRY!). Townsendia incana Nutt. var. prolixa Jones Contr. W. Bot. 13: 15. 1910. Asteraceae. = T. strigosa Nutt. Duchesne Co., Duchesne Valley, Jones 5323, 1908 (POM!;BRY!). Townsendia mensana Jones Contr. W. Bot. 13: 15. 1910. Asteraceae. Duchesne Co., near Theodore, Jones sn, 1908 (POM!;BRY!). Townsendia mensana Jones var. jonesii Beaman Contr. Gray Herb. II, 183: 88. 1957. Asteraceae. = T. jo- nesii (Beaman) Reveal Juab Co., Mammoth, Jones sn, 1910(POM!;BRY!). Townsendia minima Eastw. Leafl. W. Bot. 1: 206. 1936. Asteraceae. Garfield Co., Bryce Cyn., Eastwood & Howell 727, 1933 (CAS!). Townsendia montana Jones Zoe 4: 262. 1893. Aste- raceae. Syn: T. dejecta A. Nels. Salt Lake Co., Alta, above Flagstaff Mine, Jones sn, 1879 (POM!). Townsendia scapigera var. ambigua Gray Proc. Amer. Acad. 16: 84. 1880. Asteraceae. = T. florifer (Hook.) Gray Wayne Co., Rabbit Valley, Ward 523, 1875 (US!). Townsendia watsonii Gray Proc. Amer. Acad. 16: 84. 1880. Asteraceae. = T. florifer (Hook.) Gray Tooele Co., Stansbury Island, Watson 520, 1869 (US!). Toxicodendron longipes Greene Leafl. Bot. Obs. & Crit. 1: 118. 1905. Anacardiaceae. = T. rydbergii (Small) Greene Sevier Co., s Glenwood, Ward 212, 1875 (US!). Trifolium andersonii Gray var. friscanum Welsh Great Basin Naturalist 38: .355. 1978. Fabaceae. Beaver Co., Grampian Hill, Peabody et al 406, 1976 (BRY!;NY!). Trifolium confusum Rydb. Bull. Torrey Bot. Club 34: 46. 1907. Fabaceae. = T. longipes var. pygmaeutn Gray Iron Co., near Cedar City, Parry 35, 1874 (NY!). Trifolium inequale Rydb. Bull. Torrey Bot. Club 34: 47. 1907. Fabaceae. = T. parryi var. montanense (Rydb.) Welsh Summit Co., Bear River Canyon, Watson 243, 1869 (NY!). 186 Great Basin Naturalist Vol. 42, No. 2 Trifolium kingii Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5; 59. 1871. Fabaceae. Summit Co., Parleys Park, Watson 239, 1869 (US!;NY!). Trifolium longipes Nutt. var. brachypus Wats. Biblio. Index N. Amer. Bot. 264. 1878. Fabaceae. = T. longipes var. pijgmaeum Gray Washington Co., near St. George, Palmer sn, 1870 (GH). Trifolium macilentum Greene Pittonia 3: 223. 1897. Fabaceae. Washington Co., (s. Utah), Palmer sn, 1877 (US!;NY!). Trifolium uintense Rydb. Bull. Torrey Bot. Club 34: 47. 1907. Fabaceae. = T. dasyphyUtim var. uintense (Rydb.) Welsh Summit Co., Uintas, Watson 241, 1869 (NY!;US!). Trifolium villiferum House Bot. Gaz. 41: 335. 1906. Fabaceae. = T. eriocephahim var. villiferum (House) Martin Washington Co., (s. Utah), Palmer 91, 1877 (US!;NY!;BRY!). Triglochin maritima L. var. dehile Jones Proc. Calif. Acad. II, 5: 722. 1895. Juncaginaceae. = T. concinna var. debile (Jones) Howell Kane Co., Johnson, Jones 5289, 1894 (US!;POM!). Valeriana puberulenta Rydb. Bull. Torrey Bot. Club .36: 697. 1909. Valerianaceae. = V. acutiloba var. pub- icarpa (Rydb.) Cronq. Piute Co., Bullion Creek, Rydberg & Carlton 7065, 1905 (NY!;US!;BRY!). Valeriana pubicarpa Rydb. Bull. Torrey Bot. Club 36: 697. 1909. Valerianaceae. = V. acutiloba var. pub- icarpa (Rydb.) Cronq. Juab Co., Mt. Nebo, Rydberg & Carlton 7717, 1905 (NY!;US!;RM!). Valeriana utahensis Gandg. Bull. Soc. Bot. France 65: 37. 1908. Valerianaceae. = V. acutiloba var. pub- icarpa (Rydb.) Cronq. Cache (?) Co., Linford sn, 1907 (?). Verbesina scaposa Jones Zoe 2: 248. 1891. Aste- raceae. = Enceliopsis nutans (Eastw.) A. Nels. Grand Co., Cisco, Jones sn, 1890 (POM!;BRY!). Viguiera soliceps Barneby Leafl. W. Bot. 10: 316. 1966. Asteraceae. = Heliomeris soliceps (Barneby) Yates Kane Co., Cottonwood Canyon, Barneby 14435, 1966 (NY!;US!;UTC!). Vilfa depauperata Torr. exHook. var. filiformis Thurb. in Wats. Rep. U.S. Geol. Explor. 40th Parallel, Bot. 5: 376. 1871. Poaceae. = Muhlenbergia filiformis (Thurb.) Rydb. Summit Co., Uinta Mts., Watson 1281, 1869 (US?).' Viola beckivithii T. & G. in Beckwith Rep. U.S. Ex- plor. & Surv. R.R. Pacific 2: 119. 1855. Violaceae. Utah, between Salt Lake and Sierra Nevada, Beckwith sn, 1854(NY!). Viola beckunthii T. & G. var. cachensis C.P. Sm. Muhlenbergia 7: 136. 1912. Violaceae. = V. beckivithii T. & G. Utali. Viola bellidifolia Greene ssp. valida Baker Madrofio 5: 223. 1940. Violaceae. = V. adunca J.E. Sm. Salt Lake Co., Brighton, Baker 8519, 19.36 (UTC!). Viola bonnevillensis Cottam Bull. Univ. Utah Biol. Ser. 3: 3. 19.39. Violaceae. = V. beckivithii T. & G. Salt Lake Co., Salt Lake City, Cottam 7067, 1937 (UT!). Viola clauseniana Baker Madrofio 4: 194. 1938. Vio- laceae. = V. nephrophylla Greene Washington Co., Zion National Park, Baker 84348, 1936 (UC). Viola mamillata Greene Leafl. Bot. Obs. & Crit. 2: 33. 1910. Violaceae. = V. adunca J.E. Sm. Uintah Co., Dyer Mine, Goodding 1202, 1902 (US!;RM!;UT!;BRY!). Viola oxysepala Greene Leafl. Bot. Obs. & Crit. 2: 34. 1910. Violaceae. = V. adunca J.E. Sm. Sanpete Co., inter Willow & Ephriam creeks, Tidestrom 2476, 1909 (NDG!). Viola tidestromii Greene Leafl. Bot. Obs. & Crit. 2: 234. 1910. Violaceae. = V. adunca J.E. Sm. Sanpete Co., Ephriam Creek, Tidestrom 1143, 1908 (NDG!). Viola utahensis Baker & Clausen Leafl. W. Bot. 5: 145. 1949. Violaceae. = V. purpurea Kellogg Cache Co., Providence Canyon, Maguire 16026, 19.37 (UTC; US!;NY!;UTC!). Whipplea utahensis Wats. Amer. Naturalist 7: 300. 1873. Saxifragaceae. = Fendlerella utahensis (Wats.) Heller Kane Co., Kanab, Thompson 243, 1872 (US!). Wyethia scabra Hook. var. attenuata W.A. Weber Amer. Midi. Naturalist 35: 425. 1946. Asteraceae. Kane Co., 10.5 mi n Kanab, Carter 1424, 1938 (WS). Wyethia scabra Hook. var. canescens W.A. Weber Amer. Midi. Naturalist .35: 425. 1946. Asteraceae. San Juan Co., s Mexican Hat, Goodman & Hitchcock 1352, 19.30 (CAS!;RM!;NY!). Wyomingia vivax A. Nels. Bot. Gaz. 56: 70. 1913. As- teraceae. = Erigeron utahensis var. sparsifolius (Eastw.) Cronq. San Juan Co., Geyser Canyon, Walker 355, 1912 (US!). Xanthocephalum petradoria Welsh & Goodrich in Welsh Brittonia 33: 301. 1981. Asteraceae. Millard Co., Canyon Mts., Goodrich 15240, 1980 (BRY!;ASU!; GH!;MO!;NY!;RM!;TEX!;UC!;US!;UT!). Xylophacos aragaloides Rydb. Bull. Torrey Bot. Club 34: 48. 1907. Fabaceae. = Astragalus amphioxys var. amphioxys Washington Co., St. George, Jones 1633, 1880 (NY!;POM!;ISC!). Xylophacos marianus Rydb. Bull. Torrey Bot. Club 52: 233. 1925. Fabaceae. = Astragalus marianus (Rydb.) Barneby Piute Co., Marysvale, Jones 5355, 1894 (NY!;US!;POM!). Xylophacos medius Rydb. Bull. Torrey Bot. Club 52: 232. 1925. Fabaceae. = Astragalus eurekensis Jones Tooele Co., Lake Point, Jones 1743, 1880 (US!; CAS!;POM!;BRY!;NY!;UT!). Xylophacos melanocalyx Rydb. Bull. Torrey Bot. Club 52; 149. 1925. Fabaceae. = Astragalus amphioxys var. amphioxys Washington Co., Beaverdam Mts., Jones 5009, 1894 (NY!). Xylophacos uintensis Rydb. Bull. Torrey Bot. Club .32: 662. 1905. Fabaceae. = Astragalus argophyllus var. argophyUus Salt Lake Co., Salt Lake V., Jones 1633b, 1880 (POM!). Xylorhiza cronquistii Welsh & Atwood in Welsh Brit- tonia .33: .302. 1981. Asteraceae. Kane Co., Horse Mt., Welsh & Welsh 12819, 1975 (BRY!). Xylorhiza glabriuscula Nutt. var. linearifolia T.J. Watson Brittonia 29: 215. 1977. Asteraceae. Grand Co., 6 mi nw Moab, Watson 679, 1971 (TEX; COLO;GH;MP;MONTU;NY!;UC). Xylorhiza lanceolata Rydb. Bull. Torrey Bot. Club 37: 146. 1910. Asteraceae. = X. tortifolia var. tortifolia Washington Co., St. George, Palmer 208, 1877 (NY!;US!). June 1982 Welsh: Utah Plant Types 187 Yucca angustissima Engelm. var. avia Reveal Inter- mountain Fl. 6; 534. 1977. Liliaceae. Piute (?) Co., Loa Pass, Jones 5639a, 1898 (US!). Yucca harrimaniae Trel. Rep. Missouri Bot. Card. 13: 59. 1902. Agavaceae. Svn: Y. harrimaniae var. gilher- tiana Trel. Carbon Co., Helper, Trelease 3233, 1889 (MO!). Yucca harrimaniae Trel. var. gilbertiana Trel. Rep. Missouri Bot. Card. 18: 225. 1907. Agavaceae. = Y. har- rimaniae Trel. Juab Co., Fish Springs or House Range, Gilbert sn. 1901 (US!). Yucca kanabensis McKelvey Yuccas S.W. U.S. 2: 122. 1947. Agavaceae. Syn: Y. angustissima var. kanabensis (McKelvey) Reveal Kane Co., between Mt. Carmel & Kanab, Mckelvey 4347a, 1934 (A). Yucca toftiae Welsh Great Basin Naturalist 34: 308. 1974. .\gavaceae. Syn: Y. angustissima var. toftiae (Welsh) Reveal San Juan Co., Lake Powell, Welsh 11935a, 1973 (BRY!;NY!). Yucca utahensis McKelvey Yuccas S.W. U.S. 2: 94. 1947. Agavaceae. Syn: Y. elata var. utahensis (McKelvev) Reveal Washington Co., Santa Clara V., McKelvey 4167, 1934 (A). Zauschneria garrettii A. Nels. Proc. Biol. Soc. Wash- ington 20: 36. 1907. Onagraceae. = Epilobium canum (Greene) Raven Salt Lake Co., Big Cottonwood Canyon, Garrett 2031. 1906 (UT!). Bibliography .\l-shehbaz, I. A. 1973. The biosystematics of the genus nehjpodiiim (Cmciferae). Contr. Gray Herb. II, 204: 3-148. .\.NDERSON, L. C. 1981. Chrysothamnus nauseosus ssp. iridis (.\steraceae): a new endemic from Utah. Great Basin Naturalist 41(3): 311-313. .Atwood, N. D. 1972. New species in the Phacelia crenu- lata group (Hvdrophyllaceae). Rhodora 74: 451-463. 1973. Two new species of Phacelia (Hydro- phyllaceae). Phytologia 26: 437-438. Bailey, L. H. 1884. Notes on Carex— I. Bot. Gaz. 9:117-122. Baker, M. S. 1938. An undescribed species of Viola from Utah. Madrono 4: 194-196. 1940. Studies in western violets— III. 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Report of U.S. Geological Survey of the Fortieth Parallel. 5: xiii-525. Tidestrom, I. 1910. Species of Aquilegia growing in Utah and in adjacent portions of Colorado, Idaho and Arizona. Amer. Midi. Naturalist 1: 165-196. 1913. A new Salicornia. Proc. Biol. Soc. Washing- ton 26: 13. 1913. Novitates florae utahensis. Proc. Biol. Soc. Washington 26: 121-122. 1914. A new Delphinium from Utah. Proc. Biol. Soc. Washington 27: 61. 1923. New or noteworthy species of plants from Utah and Nevada. Proc. Biol. Soc. Washington 36: 181-184. Torrey, J. 1852. Catalogue of plants collected in the ex- pedition. Pages 381-397 in H. Stansbury, Explor- ation and survey of the valley of the Great Salt Lake of Utah. Philadelphia. Lippincott, Grambo and Co. 1871. Species of Fraxinus, and Phacelia. Pages 251,253,283. in S. Watson. Botany. Report of U.S. Geological Survey of the Fortieth Parallel. 5: xiii-525. Torrey, J., and J. C. Fremont. 1845. Pages 311-320 hi J. C. Fremont, Descriptions of some new genera and species of plants, collected in Capt. J. C. Fremont's exploring expedition to Oregon and north California in the years 1843-44. House Rep. Doc. 199. Torrey, J., and A. Gray. 1855. Report on the botanv of the expedition. Pages 115-132 in E. G. Beckwith, Report of explorations for a route for the Pacific Railroad, on the line of the forty-first parallel of north latitude. Reports of explorations and sur- veys, to ascertain the most practicable and eco- nomical route for a railroad from the Mississippi Roiover to the Pacific Ocean. 334d Cong., 2nd Sess., House Rep. Ex. Doc. Vol. II, no 91. 1870. A revision of the Eriogonea. Proc. Amer. Acad. 8: 145-200. Trelease, W. O. 1902. The Yucceae. Rep. Missouri Bot. Card. 13: 27-133. 1907. Additions to the genus Yucca. Ann. Mis- souri Bot. Card. 18: 225-230. TuzsoN, J. 1921. Arahis hirsuta f. laeiis. Bericht Freien Verein. Pflanzengeogr. Syst. Bot. 1919: 43. 1921. Underwood, L. M. 1898. Selaginella rupestris and its al- lies. Bull. Torrey Bot. Club 25: 125-133. Vasey, G. 1876. A catalogue of the forest trees of the United States which usually attain a height of six- teen feet or more. Washington. Govt. Printing Office. 1893. Descriptions of new or noteworthy grasses from the United States. Contr. U.S. Natl. Herb. 1: 267-281. Voss, J. W. 1937. A revision of the Phacelia crenulata group for North America. Bull. Torrey Bot. Club 64: 133-144. Wagner, W. L. 1981. Oenothera acutissima (Ona- graceae), a new species from northwestern Colo- rado and adjacent Utah. Systematic Botanv 6: 153-158. Ward, L. F. 1876. General notes. I— botanv. Amer. Nat- uralist 10: 552-555. Watson, E. E. 1929. Contributions to a monograph of the genus Helianthus. Papers Michigan Acad. Sci. 9: 305-475. Watson, S. 1871. Botany. In C. King. Report of the Geological Exploration of the Fortieth Parallel. 5: xiii-525. Washington: Government Printing Office. 1871. Descriptions of new species of plants, with revisions of certain genera. Proc. Amer. Acad. 12: 246-278. June 1982 Welsh: Utah Plant Types 195 1873. Revision of the extra-tropical North Ameri- can species of the genus Lupintis. Proc. Ainer. Acad. 8: 517-548. 1873. Revision of the extra-tropical North Ameri- can species of the genus PotcntiUa. Proc. .4mer. Acad. 8: 549-573. 1873. New plants of northern Arizona and the re- gion adjacent. Amer. Naturalist 7; 299-303. 1873. Revision of the extra-tropical North Ameri- can species of the genus Oenothera. Proc. Amer. Acad. 8: 573-618. 1874. A revision of the North American Che- nopodiaceae. Proc. Amer. Acad. 9: 82-126. 1874. Contributions to the botany of North America— characters of various new species. Proc. Amer. Acad. 10: 68-78. 1875. Revision of the genus oCeanothus, and de- scription of new plants. Proc. Amer. Acad. 10: 333-350. 1876. Botanical contributions— V^I. Proc. Amer. Acad. 11: 105-148. 1876. Geological Survev of California. Botanv. Vol. 1. ed. 1. 1877. Descriptions of new species of plants, with revisions of certain genera. Proc. Amer. Acad. 21: 246-278. 1878. Bibliographical index to North American botany. Washington, D.C. Smithsonian In- stitution. 476 pp. 1879. Contributions to American botanv. Proc. Amer. Acad. 14: 213-303. 1888. Contributions to American botanv. Proc. Amer. Acad. 23: 249-287. Watson, T. J., Jr. 1977. The taxonomy of Xijlorhiza (As- teraceae— Astereae). Brittonia 29: 199-216. Weber, W. A. 1946. A taxonomic and cytological study of the genus Wijethia, family Compositae, with notes on the related genus Balsomorhiza. Amer. Midi. Naturalist .35: 400-452. Welsh, S. L. 1970. An undescribed species of Astragalus (Legimiinosae) from Utali. Rhodora 72: 189-193. 1970. New and unusual plants from Utah. Great Basin Naturalist .30: 16-22. 1971. Description of a new species of Dalea (Le- giuninosae) from Utah. Great Basin Naturalist 31: 90-92. 1974. Utah plant novelties in Astragalus and Yucca. Great Basin Naturalist .34: .305-310. 1976. Utali plant novelties in Cymopterus and Penstemon. Great Basin Naturalist 35: 377-378. 1978. Endangered and threatened plants of Utah: A reevaluation. Great Basin Naturalist 38: 1-18. 1978. Utah flora: Fabaceae (Leguminosae). Great Basin Naturalist 38: 225-.367. 1980. Utah flora: Malvaceae. Great Basin Natu- ralist 40: 27-37. 1981. New taxa of western plants— in tribute. Brittonia .33: 294-.303. 1982. Utah Flora: Rosaceae. Great Basin Natural- ist (in preparation). Welsh, S. L., and N. D. Atwood. 1977. An undescribed species of Thehjpodiopsis (Brassicaceae) from the Uinta Basin, Utah. Great Basin Naturalist 37: 95-96. Welsh, S. L., and R. C. Barneby. 1981. Astragalus len- tiginosus (Fabaceae) revisited— a unique new va- riety. Isleya 2: 1-2. 1981. Welsh, S. L., and S. Goodrich. 1980. Miscellaneous plant novelties from Alaska, Nevada, and Utah. Great Basin Naturalist 40: 78-88. Welsh, S. L., and G. Moore. 1968. Plants of Natural Bridges National Monument. Proc. Utah Acad. 45: 220-248. Welsh, S. L., and J. L. Reveal. 1968. A new species of Townsendia (Compositae) from Utah. Brittonia 20: 375-377. 1977. Utah Flora: Brassicaceae (Cruciferae). Great Basin Naturalist 37: 279-365. 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 Naturali.st .35: 327-374. Wheeler, G. M. 1878. Report upon United States geo- graphical surveys west of the one hundredth me- ridian. Vol. VI. Washington D.C. Government Printing Office. 404 pp. Wherry, E. T. 1943. Microsteris, Phlox, and an inter- mediate. Brittonia 5: 60-63. 1944. New Phloxes from the Rocky Mountains and neighboring regions. Not. Nat. 146: 1-11. White, T. G. 1894. A preliminary revision of the genus Lathijrus in North and Central America. Bull. Torrey Bot. Club 21: 444-458. Williams, L. O. 1932. Field and herbarium studies— I. Bull. Torrey Bot. Club 59: 427-429. 1936. Revision of the western Primulas. Amer. Midi. Naturalist 17: 741-748. Williams, T. A. 1899. Poa fendleriana and its allies. U.S.D.A. Agrostol. Circ. 10. 6 pp. YuNCKER, T. G. 1960. Two new species of Cuscuta from North America. Brittonia 12: 38-40. A NEW SPECIES OF CRYPTANTHA (BORAGINACEAE) FROM NEVADA Kaye H. Thorne' and Larry C. Higgins- Abstract.— Described as new is Cryptantha wekhii Thorne & Higgins from the White River Valley of Nye Coun- ty, Nevada. During investigations of the flora of south- em Nevada it became evident that a pre- viously undescribed species of Cryptantha was present among the mound-forming in- habitants of the white tuffaceous deposits in the White River Valley in Nye County, Ne- vada. This silicious material is present as abandoned playa deposits along the margin of the White River Valley. The deposits sup- port a very low density of plant cover, but the species are unique. Included among the inhabitants are Frasera gypsicola, F. albo- marginata, Lepidium nana, Artemisia pyg- maea. Phlox tumulosa, and Leptodactylon caespitosum. There are other low, caespitose plants among the assemblage, but they have broader distributions. Tlie species of Cryptantha is described as follows: Cryptantha welshii Thorne & Higgins, sp. nov. Species nova Cryptantha hoffmannii I. M. Johnston proxima a qua imprimis differt foliis pustulatis inferioribus nuculis brevioribus stylo superatis 1-2 mm plantis parvioribus et tenellis. Caespitose perennial, 0.5-1.4 dm tall; stems several, erect, arising from a branched caudex, strigose to sericeus, underhair with spreading setae; leaves spatulate to oblan- ceolate, strigose or subtomentose, setose, those of the lower surface pustulate, gradu- ally reduced upwards; inflorescence cylindri- cal, densely setose, strigose, uppermost cymes elongating at maturity; calyx segments lan- ceolate, densely setose, in anthesis 3-4 mm long, in fruit becoming 5-8 mm long; corolla white, fomices yellow, 0.5 mm long, the tube shorter than the calyx, 2.5-3 mm long, 4-5 mm wide; nutlets 4, or 1-3 by abortion, broadly pear shaped to somewhat lanceolate- ovate, 1.5-2.5 mm long, 1.5-2 mm wide, margins more or less in contact, dorsal sur- face muricate to tuberculate, with a broad rugose central ridge, ventral surface scarcely muricate, the scar shortly open, triangular, the margin not elevated; style 1-1.5 mm longer than the nutlets. Type.— USA. Nevada: Nye County, White River Valley, 2.1 miles W of Sunnyside, on the road to Hot Creek Campground, T7N, R61E, S36, at 5150 ft (1567 m) elevation, ex- posed rounded ancient playa remnant of white "tuffaceous" material, occasionally mixed with sand and valley fill, 5 June 1979, K. H. Thorne and B. F. Harrison 578 (Holo- type BRY; Isotypes WTU, NY). Additional specimens: Nevada, Nye County, T7N, R62E, S31, White River Val- ley, Sunnyside Hot Creek Campground road, 2.5 miles W of Hwy 318, 5230 ft elev., salt desert shrub community, 30 June 1980, B. T. Welsh .& K. H. Thome 391 (BRY); Garden Valley, 18.4 mi S of Hot Creek Campground, in a Chrysothamnus viscidiflorus- Artemisia community, on whitish sandy clay slopes, 10 June 1980, B. F. Harrison & K. H. Thorne 13298 (BRY); White River Valley, 6.7 mi N of Hot Creek Campground, T7N, R61E, S17, on limestone hill in valley center, on white 'Herbarium, Life Science Museum, Brigham Young University, Provo, Utah 84602. 'Herbarium, Department of Biology, West Texas State University, Canyon, Texas 79015. 196 June 1982 Thorne, Higgins: a New Cryptantha 197 ^^^'-'■ Fig. 1. Cryptantha ivehhii Thorne & Higgins: A, habitat of plant; B, seed (left to right) dorsal, side, and ventral views. 198 Great Basin Naturalist Vol. 42, No. 2 outcrops, 31 May 1980, K. H. Thome & B. T. Welsh 897 (BRY). White Pine County, White River Valley, Jakes Wash, T15N, R60E, S24, pinyon-juniper and desert shrub communities with mound-forming plants on white tuf- faceous outcrops, 5 June 1980, K. H. Thome et al. 987 (BRY). Relationships of Cryptantha welshii appar- ently lie with C. hoffmannii Jtn. (Munz 1968). They are compared below: Leaves pustulate on both surfaces; nutlets ovate 3-3.5 mm long, 2-2.5 mm wide; style exceeding nutlets 0.2-0.8 mm; plant robust C hoffmannii Leaves pustulate on lower surface only; nutlets 1.5-2.5 mm long, 1.5-2 mm wide; style exceeding nutlets 1-2 mm; plant smaller, more delicate C. welshii The known distributional area of C. hoff- mannii is in the White Mountains and West- guard Pass areas of Inyo County, California (Higgins 1971). The species is named in honor of Stanley L. Welsh, whose dedication to the study of the flora of the Intermountain and Great Ba- sin regions has given a vast fund of new infor- mation concerning plant communities and rare or unusual plant species found in these regions. His help and encouragement are deeply appreciated. Literature Cited Higgins, L. C. 1971. Revision of Cryptantha subgenus Oreocarya. BYU Sci. Bull. Biol. Ser. 13(4): 1-63. Munz, P. A. 1968. A California flora and supplement. Univ. of California Press, Berkeley, California. NEW TAXA OF THISTLES {CIRSIUM; ASTERACEAE) IN UTAH Stanley L. Welsh' .\bstract.— Described are several new taxa of the genus Cirsium that occur in the state of Utah: C. eatonii (Gray) Robins, var. Iiarrisonii Welsh; C. eatonii var. iinirdockii Welsh; C. ownbetji Welsh; C. scahosum Nutt. var. thorncae; and C. virgincnsis Welsh. Diagnostic criteria are, and have been, Tlie tliistles of Utah have long constituted ^^^^^ ^^^ ^^.^^^^.^^ ^^,^^ ^^^ inconstant. Fea- one of the most difficult problems m the tures of the involucral bracts have been wide- plant taxonomy of the state. Uitterences be- , j , j- .• • i ^ ..i c f . r. 1 J 1 • 4. ly used to distmguish taxa m the group. Some tween taxa are often obscured by mter- ■' ° i i . specific hvbridization. Hybridization is not of t^ose features are better than others, m- onlv between closely related entities, but oc- eluding shape and dorsal surface texture. But curs between species that have been placed others, including degree of development of in different sections of the genus. The prob- the glandular dorsal crest and the presence of lems of interpretation are compounded by pubescence on the dorsal surface, have not tlie lack of a modern comprehensive treat- proved reliable. ment of our North American species. Pre- The following taxa have been distinguished vious treatments were based on few speci- ^j^ile attempting to provide a treatment of mens and could not account for the ^j^^ ^^,^^ ^j^.^^j^^ ^j^^ problems of our Utah variability as perceived by contemporary ^ • i r. ^ j i i u ^ ^ j t , V. 11 5 1 ^11^ • J i.1. • materials often extend beyond the state, and 1 workers. Collectors have tended to avoid this , , , . . i r .i genus and its relatives because of their spin- ^^ve had to examine materials from the sur- escent and bulky nature. Onlv dedicated per- rounding states. The entire treatment will ap- sons will trouble themselves' with the speci- pear as a portion of the paper dealing with mens, which remain as problems even after the Asteraceae (Compositae), which is in they are deposited in herbaria. The pappus progress. tends to expand, even in dry specimens, and Cirsium eatonii (Gray) Robins. Eaton tlie parachutelike apparatus floats about the Thistle. [Carduus eatonii Gray]. Three more herbarium every time someone moves the qj. \q^^ distinctive varieties are present, specimens from case to case or elsewhere. 1. Involucral bracts copiously gray- to brown- villous with long multicellular hairs; corollas ocroleucous; plants of the Uinta Mts. 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 Mts., and elsewhere 2 2(1). Involucral bracts commonly suffused with dark purple; involucres not ob- scured by outer spinose bracts; plants of the Tushar Mts 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 'Life Science Museum and Department of Botanv and Range Science, Brigham Young University, Provo, Utah 84602. 199 200 Great Basin Naturalist Vol. 42, No. 2 Var. harrisonii Welsh var. nov. Similis C. eatonii var. eatonii sed bracteis atropurpureis suffusis et spinis paucioribus. Type.— USA. Utah: Piute Co., Tushar Mts., T28S, R4W, S8, alpine meadow, talus slope, igneous gravel, 3416 m elev., 16 Aug. 1978, Welsh & Henroid 18084 (Holotype BRY). Additional specimens: Utah. Piute Co., Tushar Mts., 9 mi due W Marysvale, T27S, R5W, S35, 3050 m elev, 27 July 1976, Welsh et al. 14030 (BRY); do, T27S, R5W, SI, 28 July 1967, Welsh et al. 14050 (BRY); do, 2989 m, 25 July 1978, Welsh et al. 17738 (BRY); do, T28S, R5W, S2, 17 Aug. 1978 (BRY). The few specimens of the Tushar Mountains phase seem to average smaller than in the type variety. They are isolated from the body of the species in the islandlike Tushar Moun- tains. The variety is named in honor of Pro- fessor Bertrand F. Harrison, teacher, collec- tor, and authority on Utah grasses. 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 imder the same number in Cottonwood Can- yon (now Salt Lake County). Lodgepole pine and spruce communities upwards into alpine timdra at 2375 to 3420 m in Duchesne, Juab, Salt Lake, Summit, Tooele, and Weber coun- ties; Nevada and Colorado. Specimens from the Deep Creek Mountains have few lateral spines on the outer bracts and approach C. clavatum in technical features. More mate- rial is needed to determine their status and relationships. Var. murdockii Welsh var. nov. A C. ea- tonii var. eatonii differt in bracteis copiose griseis ad bruneis villosis pilis multicellulosis. Type.— USA. Utah: Duchesne Co.; Uinta Mts., Yellowstone Canyon, T4N, R5W, S25, 3355 m elev., Precambrian quartzite, 2 Aug. 1980, Welsh & Neese 19935 (Holotype BRY). Additional specimens: Utah. Duchesne Co.; Uinta Mts., Chepeta Lake vicinity, T5N, RIW, 828,29,32,33, spruce-lodgepole pine forest, 3233 m elev., 3 Aug 1980, Neese & Welsh 9455 (BRY); Chain Lake Basin, T4N, R4W, S23, 3377 m elev., 16 July 1979, Welsh et al. 19048 (BRY). Uintah Co., Uinta Mts., Leidy Peak, TIS, R19E, S6, 3660 m elev., Neese & Peterson 6395 (BRY); do, 30 July 1971, Waite 299 (BRY); do. White Rocks drainage, 17 July 1976, Goodrich 6478 (BRY). The plants grow in talus and rock stripes at 3230 to 3660 m in Daggett, Duchesne, and Uintah counties; endemic. The variety is named in honor of Professor Joseph Richard (Dick) Murdock, teacher and collector, whose ecological work on Oke Doke (Fifth Chain Lake) in the Uinta Mountains is clas- sic. This variety has been regarded as con- stituting a portion of C. tweedyi (Rydb.) Pet- rak. That entity was reviewed by Moore and Frankton (1965) and was mapped to include northeastern Utah in its range. No specimens were cited from Utah, however. I have seen the type of that taxon, and other material within its range in northwest Wyoming, and they differ in pubescence of involucral bracts being merely white tomentose along the margins. Cirsium ownbeyi Welsh sp. nov. 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 midrib; cauline leaves with vesture and lobing like the basal; stems 5-7 dm tall, winged-decurrent, sparingly to- mentose; involucres 1.8-2.5 cm high, 1.5-2.5 cm wide, the outermost bracts more or less pinnately spinose, lance-attenuate, smooth medially, the dorsal ridge not well devel- oped, not scabrous, sparingly tomentose along margins, the inner more or less con- torted apically; spines 3-8 mm long; corollas rose-pink. Similis Circio clavato sed in caulibus alatis et foliis tripinnatifidis differt. Type.— USA. Utah: Uintah County, Horse Trail Canyon, T4S, R24E, S4, juniper-sagebrush commu- nity, 1678 m elev., 2 July 1955, Welsh 343 (Holotype BRY). Additional specimens: Utah. Daggett County, Grouse Canyon, TIN, R25E, S2, 15 June 1978, Neese 5673 (BRY). The Ownbey thistle is known from juniper, sagebrush, and riparian communities at 1678 to 1891 m in Daggett and Uintah counties. The species is named in honor of Dr. Gerald B. Ownbey, specialist in Cirsium, who first June 1982 Welsh: New Cirsium 201 recognized the distinctive nature of this spe- cies. Relationships of the Ownbey thistle ap- parently lie with C. eatonii. Cirsium scariosum Nutt. Meadow Thistle. [Carduus lacerus Rydb., type from near Mid- way; Carduus olicescens 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.]. Our specimens fall into two rather distinctive varieties. 1. Heads 25-35 mm high, 35-80 mm wide; inner bracts slender, sometimes contorted, 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 contorted, sometimes fimbriate; plants 0-6 dm tall C. scariosum var. scariosum. Var. scariosum [Cirsium acaule var. ameri- canum Gray]. This taxon, as here interpreted, consists of an amazingly diverse assemblage that has passed imder a series of names in- cluding those cited above. Saline seeps and salt marshes, streamsides, terraces, and other meadowlands at 1310 to 3175 m in Carbon, Duchesne, Emery, Garfield, Juab, Millard, Salt Lake, Sanpete, Sevier, Summit, Tooele, and Utah counties; British Columbia to Mon- tana, south to California, Arizona, and Colo- rado. Tliis phase of C. scariosum has passed under the names C acaulescens (Gray) Schum., C. coloradoense (Rydb.) Cockerell; C. tioganum (Congdon) Petrak, C. drummondii T. & G., and C. foliosum. Nomenclature is still unclear, and more work is indicated. Our highly variable material is transitional from acaulescent to caulescent within populations, with stems, when present, that are fleshy and edible. This is our common thistle of mead- owlands, and it is unfortunate that nomencla- tural entanglements have not allowed selec- tion of an unequivocal name. Var. thorneae Welsh var. nov. A C. sca- rioso var. scarioso in capitulis majoribus et caulibus longioribus differt. Type.— USA. Utah: Piute Co., margin of Otter Creek, Grass Valley, 6.5 mi n of Angle, T28S, RIW, S30, Volcanic sediments, 2013 m elev., 11 Aug. 1976. S. L. & S. L. Welsh 14369 (Holo- type BRY). Additional specimens: Utah. Bea- ver Co., Needle Range, Vances Spring, T28S, R18W, S17, 7 July 1976, Welsh & Holmgren 13903 (BRY); do,'Wah Wah Spring, 28 Aug. 1980, Welsh et al. 20157. Garfleld Co., Pine Lake, 27 July 1977. Neese & White 3830 (BRY). Iron Co., Cedar Mt., near Navajo Lake, 28 July 1971, Higgins 4669 (BRY). Kane Co., along Skutumpa Creek, T40S, R41/2W, S76, 4 Aug. 1976, Welsh et al. 14250 (BRY). Millard Co., Pavant Mts., T21S, R21/2W, S2, 15 Aug. 1978, Welsh & Hen- roid 18031 (BRY). Piute Co., 1 mi E of King- ston, 25 July 1964, Welsh & Moore 3352 (BRY); do, 2 mi W Kingston, 20 Aug. 1976, Welsh & Taylor 14443 (BRY). In addition to the features noted above, the cauline leaves are thick, with coarse veins, and spines 8-35 mm long; endemic. Cirsium virginensis Welsh sp. nov. 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 de- current 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 spiney bracts up- wards; involucres 13-20 mm tall, 12-32 mm wide, the bracts ovate-lanceolate to narrowly lanceolate, brownish to straw colored, or of- ten suffused with purple, tomentose mar- ginally (or overall), the outer not especially reflexed, the inner serrulate or entire, smooth medially, the glandular dorsal ridge more or less developed, the apical portions of the in- ner often contorted; spines 2-6 (8) mm long, yellowish; corollas pink to lavender (or white?). Saline seems and stream terraces at 850 to 950 m in Washington Co.; Arizona. The small heads and long decurrent leaf bases are diagnostic. Ab Circio undulato distinguibili in foliis non lobato et alato decurrenti et capitulis parvioribus. Type.— USA. Utah: Washington Co., St. George, T42S, R15W, S19, hanging garden in sandstone cliffs, at 900 m elev, 13 June 1982, S. L. Welsh 21234 (Holotype 202 Great Basin Naturalist Vol. 42, No. 2 BRY; Isotypes NY, CAS, ISC, MO, POM, UT, UTC, RM). Additional specimens: Utah. Washington Co., St. George, 16 Sept. 1935, Galway 8470 (BRY;US); do, 30 June 1947, Galway 2214G (US); do, 3 June 1938, Galway sn (BRY); do, Higgins 17 Aug. 1947, Higgins 10998 (BRY). Washington Co., Beaverdam Mts., at jet of Hwy 91, and Gunlock Road, in field north of the road, T41S, R17W, S28, at ca 975 m elev, 19 Aug. 1966, Higgins 836. Arizona: Mohave Co., 1-15 river bridge, near Uttlefield, 27 Aug. 1980, Bundy 200 (BRY). Although this handsome thistle is compared to C. undulatum in the diagnosis, its relation- ships are unknown. It does not appear to be closely related to other species in the com- plex groups represented in our area. Literature Cited Moore, R. J., and C. Frankton. 1965. Cytotaxonomy of Cirsium hookerianum and related species. Canad. J. Bot. 43: 597-613. Petrak, F. 1917. Die nordamerikanischen Arten der Gattung Cirsium. Beih. Bot. Centralbl. (Abt. 2), 35: 223-567. A SPECIES OF CRYPTANTHA (BORAGINACEAE) DEDICATED TO THE MEMORY OF F. CREUTZFELDT Stanley L. Welsh' Abstract.— Described as new is Cryptantha creutzfeldtii Welsh, named to honor the memory of F. Creiitzfeldt, botanist with the Gunnison Expedition of 1853-54, who was killed along with Gunnison and other members of the party in an Indian ambush. During preparation of a manuscript deal- ing with Utah plant types and their collectors and authors, it became apparent that one of our earliest botanists has not received the at- tention of the botanical public. That person is known simply as F. Creutzfeldt, and little information is known about him. He was the botanist with the ill-fated Gunnison expedi- tion of 1853-54. That expedition was in- volved with exploration of a feasible route for a railroad from the Mississippi River to the Pacific Ocean (Beckwith 1854). Creutzfeldt died (on 26 October 1853) along with Gunnison and other members of the party in an Indian attack while they were camped along the Sevier River, near present- day Delta. Captain John Williams Gunnison was in command of the expedition, which had been split prior to the Indian attack. Lieutenant Edward Griffin Beckwith was in charge of a portion of the expedition that was working separately from Gunnison at the time of the disaster. Beckwith was named to succeed Gunnison. Creutzfeldt had collected types of Eriogonum corymbosiim Benth. in DC. var. divaricatiim T. & G. in Beckwith and E. leptocladon T. & G. in Beckwith in the vicinity of the Green River crossing. The specimens of those taxa are deposited at NY. Plant taxa have been named after both Gun- nison and Beckwith, but none have been named in honor of the memory of the one person who gave more than anyone to the cause of plant collection in Utah— his life. Therefore, it is with humility and a sense of gratitude that I name the following plant after F. Creutzfeldt, who passed to the west. south of where this plant has existed undes- cribed since his time. Cryptantha creutzfeldtii Welsh, sp. nov. Similis Cryptantha jonesiana (Payson) Pay- son sed in folius longioribus et acutis glabris supra et acutis vel obtusis staturis elatioribus et calycibus majoribus differt. Perennial herbs, 0.7-2.3 dm tall; stems many, arising from a multicipital caudex and stout black-barked taproot, the caudex branches 2-12 cm long clothed by marces- cent leaf bases; leaves narrowly spatulate to oblanceolate, acute to obtuse, 2-8 cm long, 0.2-0.9 cm wide, coarsely appressed setose- pustulate (appearing ashy gray on leaves of previous years) below, the petioles long-se- tose; inflorescence an interrupted thyrse, with few to several clusters below the termi- nal subcapitate one; calyx segments lance-lin- ear in anthesis, 6-8 mm long, in fruit 9-13 mm long, densely long-setose with yellowish, ascending bristles; corolla white, the tube 8-11 mm long, campanulate in the throat; the fornices low and broad, crests at base of tube lacking, the limb 10-13 mm wide; nut- lets lanceolate, 4-5 mm long, muricate; scar narrow, open, without an elevated margin. Type.- USA. Utah: Emery County, T21S, R6E, S22, ca 1.5 mi nw hwy 10, along dirt road at Muddy Creek historical marker, on slope S of canal, 6400 ft (1952 m), shadscale community, 14 May 1981, S. L. Welsh 20470 (Holotype BRY; isotypes NY, ISC, POM, CAS, MINN, COLO, RM, UT, UTC, UC, MO, and others). 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 203 204 Great Basin Naturalist Vol. 42, No. 2 Additional specimens: Utah, Carbon Co., T14S, RIOE, S17-18, ca 1 mi W Price, Va mi S Castleview Hospital, Mancos Shale slope at ca 5500 ft (1678 m) elev., 19 May 1981, L. Arnold sn (BRY); NY, ISC, POM, CAS, MINN, COLO, RM, UT, UTC, UC, MO, RENO. Emery Co., T21S, R6E, S27, 5 mi SW of Muddy Creek, 3 mi N Emery, Bluegate Member, clay soil, 26 May 1979, E. Neese & S. White 7.362 (BRY); Castle Dale, SE Buz- zard Bench, 20 May 1976, J. Allan 762, (BRY); T22S, R6E, NEV4 S7, 1.7 mi due W Emery; 30 May 1979, S. White 15 (BRY); T21S, R6E, S28, 1.5 mi due N Emery, 7 June 1979, S. White & G. Moore 77 (BRY); T20S, R6E, S14, Ferron Canyon, 23 May 1980, N. D. Atwood 7518 (BRY); SW of Orangeville, below Buzzard Bench, 27 April 1979, J. Allan 947 (BRY). The Creutzfeldt cryptantha is a near ally of Cryptantha jonesiana (Payson) Payson. Di- agnostic features of the former include the acute to obtuse leaves that are glabrous above, the stature that averages taller at maturity, the larger calyces, and the large nutlets. The two taxa are spatially and edaphically isolated, with C. jonesiana oc- cupying portions of the Summerville and Moenkopi formations in the central portions of the San Rafael Swell, and C. creutzfeldtii occurring on the Blue Gate Member of the Mancos Shale Formation along the strike of that formation below the coal measures of the Mesa Verde Group of formations. Both flower in early springtime, and each is char- acterized by the broad corolla limbs, vesture of pustulate-based setae, and slender accres- cent calyx lobes. Literature Cited HiGGiNS, L. C. 1971. A revision of Cryptantha subgenus Oreocarya. BYU Sci. Bull., Biol. Ser. 13 (4). Beckwith, E. G. 1855. Reports of explorations and sur- veys, to ascertain the most practicable and eco- nomical route for a railroad from the Mississippi River to the Pacific Ocean. House Rep. Exec. Doc, Vol. 2, No. 91. ALGAL POPULATIONS IN BOTTLE HOLLOW RESERVOIR, DUCHESNE COUNTY, UTAH Jeffrey Johansen', Samuel R. Rushforth-, and Irena Kaczmarska' Abstract.— Bottle Hollow Reservoir contains a diverse algal flora. A total of 289 taxa was observed, 227 of which were diatoms. Both littoral and planktonic communities had high diatom diversity. During summer months fila- mentous Chlorophvta were diverse and high in biomass in the littoral zone. Phytoplankton collections in Bottle Hol- low Reservoir were dominated by four species: Asterionella formosa, Cyclotella comta, Dinobryon dwergens, and Fragikiria crotonensis. Plankton samples contained mostly small diatoms in early spring, with larger algae succeeding the.se as the summer progressed. No blue-green algae were important in this succession. Two peak production peri- ods were observed, one in the fall and one in the spring. Bottle Hollow Reservoir appears to be a healthy mesotro- phic svstem based on the evidences of moderately high algal diversity, insignificance of blue-green algae, and the presence of a suite of diatom species indicative of mesotrophic conditions. Bottle Hollow Reservoir is in Fort Duch- esne, Duchesne County, Utah, on the Ute In- dian Reservation. It was planned and con- structed by the Bureau of Reclamation as a mitigation component of the Bonneville Unit of the Central Utah Project. The primary function of this reservoir was to replace part of the fisheries and recreation lost due to the construction of the Rock Creek component of the Central Utah Project. Bottle Hollow Reservoir is presently the central component of tlie Bottle Hollow Resort owned and oper- ated by the Ute Indian Tribe. It is used pri- marily for sport fishing. Construction of Bottle Hollow Reservoir was completed in 1971 and the lake was filled during 1972. Water for the reservoir is taken from the Uinta River through the In- dian Bench Canal that originates 11 km to the northwest. Little outflow is released from the reservoir at any time of the year. Water to replace that lost by evaporation and seep- age is brought from the Uinta River through the Indian Bench Canal during the early spring. No appreciable flushing or flow- through has occurred in the reservoir since its completion. Total capacity of Bottle Hollow Reservoir is 11,103 acre feet, with usable ca- pacity at essentially the same figure. Eleva- tion of the spillway is 1552.8 meters, and sur- face area of the reservoir is 418 acres. Fishing in Bottle Hollow has been good to excellent since its completion. The fishery is based primarily upon planted brown trout (Merritt et al. 1980). Concern to maintain this fishery and concern over somewhat poorer catches during the past few years led the Ute Indian Tribe to initiate a com- prehensive study of the water quality and bi- ology of the system. This study was financed by the Environmental Protection Agency through an areawide 208 water quality plan- ning grant. We have studied the algal floras of this reservoir during 1977 and 1979-1980. Methods Ten collection stations were established to monitor the plankton and attached algae in Bottle Hollow Reservoir (Fig. 1). The first five were identical to the water quality sta- tions used by Merritt et al. (1980) for chem- ical and physical analyses of the reservoir. Littoral collections were made at four sites around the periphery of the reservoir: the shore near the inlet channel, the north end, the south dam, and the south end. The inlet channel itself was also sampled. Four series 'Department of Oceanography, Texas A & M University, College Station, Texas 77843. -Department of Botany, Brigfiam Young University, Provo, Utah 84602. 'Polish Academy of Sciences, Institute of Botany, Department of Phycology, ul. Lubicz 46, 31-512 Krakow, Poland. 205 206 Great Basin Naturalist Vol. 42, No. 2 of collections were made during the 1979-80 study period (Table 1). Plankton samples were collected using a 2.3 liter capacity Van Doren bottle. Four Van Doren bottles distributed evenly through the euphotic zone were collected at each phytoplankton site and filtered through a 35 mm mesh phytoplankton net into a large bucket, yielding a composite net plankton sample for each site. In addition, a composite nannoplankton sample was collected by sam- pling the filtrate in the bucket. Sediment samples were collected using an Ekmann Dredge. Littoral algal collections were chiefly of attached species, though twice unattached filamentous green algae were collected (Table 1). Attached algae consisted of epiphyton (algae growing on vascular plants), epilithon (algae attached to rocks), and epip- sammon (algae growing on and in sand or silt). Samples were returned to Brigham Young University and placed under refrigeration. Analyses of living algae were made within one week after collection. Nannoplankton V " ( Boat Launch Area y \ r J Indian Canal y' \ Inlet Channel j \^ North Dam \ ■' [ V 2.7 \ South Dam Bottle Hollow Reservoir { !X 1 Vec.ty vernal Provo V Bottle Hollow/ Reservoir UTAH 1 1 Fig. 1. Reference map of Bottle Hollow Reservoir showing the collecting localities. samples were concentrated by vacuum filtra- tion through Millipore filters (1.2 mm pore size). Estimates of absolute densities of plank- tonic algae were made using Palmer Cell wa- ter mounts. Living algae in littoral and ben- thic samples were identified and the abundance of each species estimated. After living algae were studied, the dia- toms in each sample were cleaned, using standard nitric acid oxidation techniques (St. Clair and Rushforth 1976), and mounted in Hyrax resin. All algae were examined and identified using Zeiss RA research micro- scopes with phase contrast and Nomarski in- terference phase accessories. Results A total of 280 algal taxa were observed during this study. Twenty-three of these were blue-green algae (Cyanophyta); 32 were green algae (Chlorophyta); 4 were eu- glenophytes (Euglenophyta); 2 were dinofla- gellates (Pyrrhophyta); one was a chryso- phyte (Chrysophyta); and 227 were diatoms (Bacillariophyta). All algal species, together with their occurrence in the major micro- habitats of the reservoir, are listed in Table 2. Living algae were not observed in any of the sediment collections, and so diatom slides made from these samples were not quan- titatively analyzed. Littoral communities were dominated by filamentous green algae most of the year. These were chiefly representatives of Table 1. Algal samples collected from Bottle Hollow Reservoir during the 1979-1980 sampling period. All samples were examined for nondiatoms. Permanent dia- tom slides of samples from stations 1-6 and 9 were also examined. Key: P = plankton; S = sediments; Ep = epiphytic algae; El = epilithic algae; Es = epipsammic algae; L = littoral unattached algae. Station 15 Nov 1979 27 Mar 1980 20 Jun 1980 26 Jul 1980 1 P, s P, s 2 P, s P, s P P 3 P, s P, s 4 P, s P, s 5 P, s P, s 6 Ep, Es Ep, Es Ep, Es El, Es 7 El, Es El El 8 El, Es, L L El, Es 9 Ep, Es Ep, Es Es 10 El El El June 1982 JOHANSEN ET AL.: AlGAL POPULATIONS 207 Table 2. Algal species collected from Bottle Hollow Reservoir, Duchesne County, Utah, with their distribution in the various habitats studied. Species a g o S ^ Oi a, a, CU W W Cyanoph\ta Anabaena variabilis Kuetzing Anabaena sp. Aphanizomenon flos-aqtiae (Lenim.) Ralfs Cahthrix sp. Chroococcus liinneticiis Lemmermann Chroococcus turgidus (Kg.) Naegeli Gloeocapsa decorticans (A. Br.) P. Richt. Gomphos'phaeria aponina var. delicatula Virieux Lijngbya birgii CM. Smith Lyngbya diguetii Gomont Merismopedia glauca (Ehr.) Naegeli Nodularia spumigena Mertens Oscillatoria agardhii Gomont Oscillatoria angusta Koppe Oscillatoria geminata Schwabe Oscillatoria limnetica Lemmermann Oscillatoria limosa (Roth) Agardh Oscillatoria subbrevis Schmidle Oscillatoria tenuis Agardh Oscillatoria sp. Phormidiiim teniie (Menegh.) Gomont Spindina major Kuetzing Tolypothrix distorta Kuetzing Chlorophyta Ankistrode sinus falcatus (Corda) Ralfs Chlamydomonas globosa Snow Cladophora glomerata (Lemm.) Kuetzing Closterium dianae Ehrenberg Cosmarium nitidulum De Not. Cosrnariiim sp. Dictyosphaerium ehrenbergianwn Neageli Eudorina elegans Ehrenberg Mougeotia sp. Oedogonium sp. 1 Oedogonium sp. 2 Oedogonium sp. .3 Oedogonium sp. 4 Oedogonium sp. 5 Oedogonium sp. 6 Oocystis gloeocystiforrnis Borge Oocystis pusilla Hansgirg Pediastrum boryanum (Turp.) Meneghini Rhizoclonium hieroghjphicum (Ag.) Kuetzing Rhizoclonium sp. Scenedesrnus bijuga (Turp.) Lagerheim Scenedesmus quadricauda var. longispina (Chod.) G.M. Smith Scenedesrnus quadricauda var. quadrispina (Turp.) Brebisson Sphaerocystis schroeteri Chodat Spirogyra sp. 1 Spirogyra sp. 2 Spirogyra sp. 3 Spirogyra sp. 4 Spirogyra sp. 5 208 Table 2 continued. Great Basin Naturalist Vol. 42, No. 2 Species a. 'a, w Oh H Oh Staurastrum gracile{?) Ralfs Ulothrix zonata (Weber et Mohr) Kuetzing Zygnema sp. EuGLENOPHYTA Euglena elastica Prescott Euglena gracilis Klebs Trachelomonas ahrupta (Swir.) Deflandre Trachelomonas dybowskii Drezepolski Pyrrhophyta Ceratiiim hirudinella (O.F. Muell.) Dujardin Glenodinium pulvisculus (Ehr.) Stein Chrysophyta Dinobryon divergens Imhof Bacillariophyta Achnanthes clevei Gmnow Achnanthes conspicua A. Mayer Achnanthes exigua Grunow Achnanthes gibberula Grunow Achnanthes haiickiana Grunow Achnanthes kryophila Petersen Achnanthes lanceolata (Breb.) Grunow Achnanthes lanceolata var. diibia Grunow Achnanthes Hnearis (W.Sni.) Grunow Achnanthes linearis f. carta H.L. Smith Achnanthes minutissima Kuetzing Achnanthes orientalis Hustedt Achnanthes peragalU var. fossihs Tempere & Peragallo Achnanthes sp. 1 Achnanthes sp. 2 Amphipleiira pellucida Kuetzing Amphora coffeiformis (Ag.) Kuetzing Amphora ovalis (Kg.) Kuetzing Amphora ovalis var. affinis (Kg.) v. Heurck ex De Toni Amphora ovalis var. pediculus (Kg.) v. Heurck ex De Toni Amphora perpusilla (Grun.) Grunow Amphora veneta Kuetzing Anomoeoneis serians (Breb. ex Kg.) Cleve Anonweoneis serians var. brachysira (Breb. ex Kg.) Hustedt Anonweoneis sphaerophora (Kg.) Pfitzer Anonweoneis zellensis (Grun.) Cleve Asterionella fomiosa Hassall Bacillaria paxillifer (O. Muell.) Hendey Biddtdphia levis Ehrenberg Caloneis amphisbaena (Bory) Cleve Caloneis bacillum (Grun.) Cleve Caloneis lewisii Patrick Caloneis lewisii var. inflata (Schultze) Patrick Caloneis ventricosa var. truncatula (Grun.) Meister Chaetoceros sp. Cocconeis pedicidus Ehrenberg Cocconeis placentula Ehrenberg June 1982 Johansen et al.: Algal Populations 209 Table 2 continued. Species a, ^ o- a- u w Cocconeis placentula var. euglypta (Ehr.) Cleve Cocconeis placentula var. lineata (Ehr.) Cleve CyclotelUi caspia Grunow Cyclotella cotnta (Ehr.) Kuetzing Cyclotella rneneghiniana Kuetzing Cyclotella sp. Cylindrotheca gracilis (Breb.) Grunow Cymatopleura solea (Breb.) W. Smith Cymbella affinis Kuetzing Cyrnbella brehmii Hustedt Cymbella cuspidata Kuetzing Cymbella cymbifonnis Agardh Cymbella mexicana (Ehr.) Cleve Cymbella microcephala Grunow Cymbella miniita Hilse ex Rabenhorst Cymbella mintita var. latens (Krasske) Reinier Cymbella miniita var. silesiaca (Bleisch ex Rabh.) Reimer Cymbella muelleri Hustedt Cymbella norvegica Gninow Cymbella siniiata Gregory Cymbella titmida (Breb.) v. Heurck Cymbella turgidula Grunow Cymbella sp. 1 Cymbella sp. 2 Denticula elegans f. valida Pedicino Denticula sp. Diatoma tenue Agardh Diatcrma tenue var. elongatum Lyngbye Diploneis oculata (Breb.) Cleve Diploneis subovalis Cleve Entomoneis ornata (Bail.) Reimer Epithemia adnata var. proboscidea (Kg.) Patrick Epithemia argus var. protracta A. Mayer Epithemia smithii Carruthers Epithemia sorex Kuetzing Epithemia turgida (Ehr.) Kuetzing Fragilaria brevistriata Grimow Fragilaria brevistriata var. inflata (Pant.) Hustedt Fragilaria cf capucina Desmazieres Fragilaria capucina var. mesolepta Rabenhorst Fragilaria construens var. venter (Ehr.) Grunow Fragilaria crotonensis Kitton Fragilaria crotonensis var. oregonica Sovereign Fragilaria leptostauron (Ehr.) Hustedt Fragilaria leptostauron var. dubia (Grun.) Hustedt Fragilaria pinnata Ehrenberg Fragilaria similis Krasske Fragilaria vaucheriae (Kg.) Peterson Fragilaria virescens Ralfs Frustulia vtdgaris (Thw.) De Toni Gomphoneina acuminatum Ehrenberg Gomphonemo' affine Kuetzing Gomphonema dichotomum Kuetzing Gomphonema instabilis Hohn & Hellerman Gomphonenui intricatum Kuetzing 210 Table 2 continued. Great Basin Naturalist Vol. 42, No. 2 Species o s S CL, a a, a W W Gomphonema olivaceum (Lyngb.) Kuetzing Gomphonenui olivaceum var. calcarea (CI.) Cleve Gomphonema parvuhim Kuetzing Gomphonema parvuhim var. micropus (Kg.) Cleve Gomphonema suhclavatum (Gnin.) Grunow Gomphonema truncatum Ehrenberg Gomphonema sp. Gyrosigma acuminatum (Kg.) Rabenhorst Gyrosigma fasciola (Ehr.) Griffith & Henfrey Gyrosigma obtusatum (Sulliv. & Wormley) Boyer Hannaea arcus (Ehr.) Patrick Hantzschia amphioxys (Ehr.) Grunow Hantzschia distincte-punctata (Hust.) Hustedt Hantzschia virgata (Roper) Grunow Mastogloia braunii Grunow Mastogloia smithii var. lacustris Grunow Melosira granulata (Ehr.) Ralfs Navicula anglica Ralfs Navicula anglica var. subsaka (Grun.) Cleve Navicula arvensis Hustedt Navicula atomus (Kg.) Gninow Navicula bacillifonnis Gnmow Navicula capitata Ehrenberg Navicula capitata var. hungarica (Grun.) Ross Navicula capitata var. lunebergensis (Grun.) Patrick Navicula cincta (Ehr.) Ralfs Navicula circumtexta Meister ex Hustedt Navicula clementoides Hustedt Navicula contenta f. biceps (Arnot.) Grunow Navicula cryptocephala Kuetzing Navicula cryptocephala var. exilis (Kg.) Grunow Navicula cryptocephala var. veneta (Kg.) Rabenhorst Navicula cuspidata (Kg.) Kuetzing Navicula decussis Oestrup Navicula disjuncta Hustedt Navicula elginensis (Greg). Ralfs Navicula exigua var. capitata Patrick Navicula gastrum Ehrenberg Navicula graciloides A. Mayer Navicula grimmei Krasske Navicula halophila (Grun.) Cleve Navicula halophila f. tenuirostris Hustedt Navicula heufleri Grunow Navicula heufleri var. leptocephala (Breb. ex Grun.) Patrick Navicula luzonensis Hustedt Navicula menisculus var. upsaliensis (Grun.) Gmnow Navicula minima Grunow Navicula mutica var. cohnii (Hilse) Grvinow Navicula mutica var. undulata (Hilse) Gmnow Navicula oblonga Kuetzing Navicula pelliculosa (Breb. ex Kg.) Hilse Navicula peregrina (Ehr.) Kuetzing Navicula permitis Hustedt Navicula pupula Kuetzing Navicula pupula var. mutata (Krasske) Hustedt June 1982 Table 2 continued. JOHANSEN ET AL.: AlGAL POPULATIONS 211 Species a CL, Navicula pupiila var. rect/jngiilaris (Greg.) Grunow Navicida pijonuiea Kuetzing Navicula radiosa Kuetzing Navicula radiosa var. tenella (Breb. ex Kg.) Gmnow Navicula diynchocephaki Kuetzing Navicula salinarum var. intermedia (Grun.) Cleve Navicula secreta var. apiculata Patrick Navicula tantida Hustedt Navicula tenelloides Hustedt Navicula tenera Hustedt Navicula tripunctata (Muehl.) Bory Navicula tripunctata var. schizonemoides (v. Heurck) Patrick Navicula viridula (Kg.) Kuetzing Navicula viridula var. linearis Hustedt Navicula viridula var. rostellata (Kg.?) Cleve Navicula sp. 1 Navicula sp. 2 Navicula sp. 3 Navicula sp. 4 Navicula sp. 5 Navicula sp. 6 Neidium bisulcatum var. baicalense (Skr. & Meyer) Reinier Neidium dubium (Ehr.) Cleve Nitzschia acicularis W. Smith Nitzschia acicularoides Hustedt Nitzschia ci amphibia Grunow Nitzschia angustata (W. Sm.) Gmnow Nitzschia apiculata (Greg.) Grunow Nitzschia circumsuta (?) (Bail.) Grunow Nitzschia dissipata (Kg.) Grunow Nitzscliia frustidurn Kuetzing Nitzschia gandersheimensis Krasske Nitzschia hantzschiana Rabenhorst Nitzschia hungarica Grunow Nitzschia inconspicua Grunow Nitzschia microcephala Grunow Nitzschia minutula Grunow Nitzschia ovalis Arnott Nitzschia palea (Kg.) W. Smith Nitzschia palcacea Grunow Nitzschia punctata (W. Sm.) Gnmow Nitzschia pusilla (Kg.) Grun. em. Lange-Bertalot Nitzschia recta Hantzsch Nitzschia romana Gnmow Nitzschia sigma var. sigmatella Gnmow Nitzschia sigmoidea (Ehr.) W. Smith Nitzschia sinuata (W. Sm.) Grunow Nitzschia sinuata var. tabellaria Gnmow Nitzschia sociabilis Hustedt Nitzschia trybionella var. debilis (Arnott) A. Mayer Nitzschia trybionella cf var. levidensis (W. Sm.) Grunow Nitzschia trybionella var. victoriae Grunow Nitzschia valdestriata Aleem & Hustedt Nitzschia sp. 1 Nitzschia sp. 2 212 Table 2 continued. Great Basin Naturalist Vol. 42, No. 2 Species Pinntilaria abaujensis var. linearis (Hust.) Patrick Pinntilario borealis Ehrenberg Pinnularia brebissonii Kuetzing Pinnularia brebissonii var. diminuta (Gnin.) Cleve. Pleurosigma sp. Rhoicosphenia curvata (Kg.) Gninow Rhopalodia gibba (Ehr.) O. Mueller Rhopahdia gibberuki (Ehr.) O. Mueller Rhopalodia gibberula var. vanhettrckii O. Mueller Staiironeis anceps Ehrenberg Stauroneis smithii Grunow Staiironeis ci smithii Grunow Stauroneis wislouchii For. et Anisini. Stephanodiscus astraea var. minutula (Kg.) Gninow Stephanodisciis niagarae Ehrenberg Stephanodiscus sp. Surirella angusta Kuetzing Surirella ovalis Brebisson Synedra acus Kuetzing Synedra cyclopum Brutschi Synedra fasciculata (Ag.) Kuetzing Synedra fasciculata var. truncata (Grev.) Patrick Synedra pulchellu Kuetzing Synedra radians Kuetzing Synedra ulna (Nitzsch) Ehrenberg Synedra ulna var. ramesi (Herib. et Perag.) Hustedt Zygnematales (Spirogyra, Mougeotia, and Zygnema species), though Oedogonium spe- cies were also important. Because sexual stages were not observed, these taxa could not be identified beyond the generic level. Cladophora glomerata was important in the inlet channel but was also occasionally com- mon in some littoral sites of the reservoir. Ulothrix zonata was abundant in the channel in November, but was only rarely observed in the reservoir. Diatoms were also important in the littoral commimities, and dominated the algal assemblage during the winter and early spring. Filamentous Chlorophyta died off during the winter and were not reestablished until early summer. Diatoms on the other hand recovered soon after winter ice had melted. The eight most important diatoms in the littoral sites were all pennate species; Di- atonia teniie, Fragilaria vaucheriae, Gompho- nema instabilis, Navicula cryptocephala var. veneta, Navicula rnutica var. cohnii, Nitz- schia minutula, Nitzschia palea, and Nitz- schia paleacea (Table 3). The three different substrata sampled showed differences in diatom floras. Coc- coneis placentula var. lineata, Cyynbella min- uta var. silesiaca, Gomphonema olivaceum, and Gomphonema instabilis were primarily epiphytes. The epipsammon was character- ized by small raphoid diatoms, particularly Navicula mutica var. cohnii. Filamentous green algae were either unattached or part of the epilithon and epiphyton. Most algal spe- cies in the littoral were at least to some de- gree cosmopolitan. Many algal species are opportunistic gen- eralists (Lowe 1974, Patrick and Reimer 1966). Achnanthes minutissima, Navicula cryptocephala var. veneta, Nitzschia palea, and Nitzschia paleacea, as well as several other small raphoid diatom species in the study, are such taxa. These organisms occur in a wide variety of habitats in western North America and worldwide (Camburn et al. 1978, Foged 1959, 1974, Hustedt 1930, 1949, Patrick and Reimer 1966). Other diatoms are June 1982 JOHANSEN ET AL.: AlGAL POPULATIONS 213 best suited to grow in more specialized env ronments. For example, many species in the genus Cocconeis grow optimally on sub- merged macrophytes (Lowe 1974, Patrick and Reimer 1966). These species can also be foimd on rocks or wood and, through mixing processes in the lake, will also occur in the epipsammon and plankton. Because species are not confined to the substrate on which they are best suited, characterizing species according to habitat preference is often diffi- cult. Even so, the planktonic algal assem- blages in Bottle Hollow Reservoir were dis- tinctly different from those of the littoral, despite the overlap of some species. The dominant algal plankters were limited to three diatom taxa and one chrysophyte; As- terionella fomiosa, Cyclotello comta, Fragi- laria crotonensis, and Dinobryon divergens (Table 3). These species usually composed about 80 percent of the total planktonic flora. Because of this, diversity was much lower in the plankton than in the periphyton. Total phytoplankton abundance ranged from an average density as low as 700,000 organ- isms per liter in late July to a high of 1,700,000 organisms per liter in November. Discussion Three areas of interest concerning the floras of Bottle Hollow Reservoir will be dis- cussed; floristic diversity, community dynam- ics, and trophic condition. It has already been noted that diversity in the planktonic envi- Table 3. Average percent densities of the 14 most important diatom taxa in Bottle Hollow Reservoir. Aver- age densities were computed separately for plankton and littoral samples. Species Plankton Littoral Achnanthes mintitissima .3 2.5 Asteriunella formosa 25.5 .4 Cyclotella comta 17.1 .7 Cymbella microcephah .2 2.2 Diatoma tenue .4 3.8 Fragihiha crotonensis 39.5 1.6 Fraphirid vauclwhae .1 3.1 Gomplioucnui instahilis .1 3.9 XaviCuki cnjptoccpliala var. veneta .4 8.4 Xavicuhi mutica var. cohnii .1 7.9 Xitzschia microcepliala .2 2.1 Xitzscliki minutula .1 3.2 Xitzschia palea 2.1 3.8 Xitzschia paleacea 2.7 4.8 ronment was depressed by the dominance of four algal taxa. Even so, a total of 174 algal taxa (154 of which were diatoms) were found in the phytoplankton samples. This is 60 spe- cies fewer than found in the littoral zone, which had a total of 234 taxa (184 of which were diatoms). The diversity in phytoplank- ton was due primarily to the infrequent oc- currence of small diatom species in the water column. These species are easily transferred from the littoral and benthic areas, where they are often most common, to the open wa- ter of the lake by natural mixing processes. A few supposed littoral species such as Ach- nanthes orientalis were more common in the plankton than in the littoral collections, but these were more the exception than the rule. Because the majority of the littoral-plankton- ic diatoms were small, they were found pri- marily in nannoplankton samples and were much less frequent in the netplankton. Net- plankton samples had an average of 16 dia- tom taxa per sample, whereas nannoplankton collections contained an average of 30 taxa (Table 4). Littoral collections contained sub- stantially greater numbers of species. This is to be expected because the littoral environ- ment is more heterogenous than the plank- tonic habitat and contains more ecological niches. The highest number of species per sample was found in the November littoral collections (Table 4). Population dynamics of the plankton are easier to monitor than those for the littoral areas. This is largely due to the relative ease of obtaining quantitative phytoplankton data versus quantitative data for attached species. If the numbers of netplankton individuals per liter are added to the numbers of nanno- plankton individuals per liter from the same locality, estimates of total phytoplankton per hter of lake water are obtained. Average den- sities of the four most abundant taxa were computed using these estimates and plotted against time (Fig. 2). Fragilaria crotonensis was the most abundant species, reaching higher concentrations as the seasons pro- gressed. The highest density of this taxon was observed in the November 1979 collections. Several observations and speculations can be made after consideration of the data shown in Figure 2. Two periods of peak algal production, fall and spring, occur in Bottle 214 Great Basin Naturalist Vol. 42, No. 2 Total Phytoplankton Four most abundant taxa Astenonella lormosa Cyclotella comta Dinobryon divergens Fragilana crotonensis Nov Dec Jan Feb Mar Apr May Jun Jul Aug Sep Fig. 2. Densities of the dominant phytoplankters and total phytoplankton through the 1979-1980 collecting year in Bottle Hollow Reservoir. Hollow Reservoir. These are likely due to fall and spring turnover. During winter, produc- tion falls drastically with shorter days and ice coverage. As soon as the ice melts, small dia- toms grow quickly in the recently mixed, nu- trient-rich water. The March collections had substantial numbers of these small diatoms, even though the biomass was still quite low. Cyclotella comta was present in higher num- bers than F. crotonensis at this time. The pulse of these small algae favors the growth of small filter-feeder zooplankton (Porter 1977), such as the cladocerans that were observed in both the March and June net hauls. As the zooplankters apply a selec- tive pressure on small diatoms, larger (often colonial) algae may become more prevalent (Porter 1977, Wimpenny 1973). The density of Cyclotella comta in Bottle Hollow Reser- voir leveled off in June and dropped drasti- cally in July. The larger colonial forms As- terionella formosa, Dinobryon divergens, and Fragilaria crotonensis increased in late spring and dominated the spring peak. As summer progressed, total phytoplank- ton density dropped, though F. crotonensis continued to increase in number. This may be due to two factors. First, the lake begins to stratify during early spring, causing mixing to cease. Nutrients tied up in the living algae and zooplankton are lost to the sediments as these organisms die and sink and as feces of zooplankton and fish settle (Wetzel 1975). Second, grazing pressure may decrease total phytoplankton density as zooplankton popu- lations reach maturity (Porter 1977). Filter- feeders cannot feed well on the large Fragi- laria colonies, and so F. crotonensis tends to escape predation and continues to increase in number. An unexplained phenomena is the decrease in the large colonial algae Asterio- nella formosa and Dinobryon divergens, which should also have the same size refuge from filter feeders as F. crotonensis. The de- cline of A. formosa in early summer is a com- mon occurrence that has been attributed to nutrient depletion in the upper water (Pear- sail 1932). Another possibility is that larger raptorian-feeder zooplankton, such as many copepods, which begin to reach maturity lat- er in summer, may have a preference for these algae over F. crotonensis. Finally, it is clear that either or both of these algae could decrease due to temperature increase or some other environmental factor. Littoral algal succession was less well de- fined. Diatoms were particularly important in early spring and grew to some extent when the lake was covered with ice. As the water warmed, filamentous green algae became im- portant and had the highest standing crop. Despite the higher biomass of these green al- gae, diatoms may be more critical to littoral Table 4. Average number of diatom species per microhabitat type Microhabitat November March June July Average Netplankton Nannoplankton Epiphyton Epipsammon 18.4 18.2 75.5 63.5 12.4 40.4 46.5 21.5 22.0 21.0 62.0 48.0 51.0 64.0 16.0 30.4 61.2 50.9 June 1982 JOHANSEN ET AL.: AlGAL POPULATIONS 215 15 16 \ 17 18 19 20 Figs. 3-20. Diatom spp.: 3, Cijclotella comta. 19 mm diameter, 12 striae/10 mm; 4, Cocconeis placentula var. line- ata, 15 X 11 mm, 19 striae/10 mm; 5, Achnanthes exigtia, 12.5 X 5 mm, 24 striae/10 mm; 6, A. orientalis. Raphe valve: 11.5 X 4 mm, 26-30 striae/10 mm; 7, A. orientalis. Rapheless valve; 12 X 4 mm, 26-30 striae/10 mm; 8, Diploneis suh- ovalis, 14 X 8.5 mm, 11 costae/10 mm; 9, Diatoma tenue, 29 X 3 mm, 7-10 costae/10 mm; 10, Navicula cnjptocephala, 32 X 6 mm, 15-18 striae/10 mm; 11, .V. salinarum var. intermedia, 35 X 7 mm, 14-16 striae/10 mm; 12, A', tripunc- tata, 32 X 6 mm, 12-14 striae/ 10 mm; 13, N. tenera, 13 X 4.5 mm, 20 striae/10 mm; 14, N. sp. 3, 10 X 5 mm, 30 striae/10 mm; 15, N. cryptocephala var. veneta (?), 22 X 5 mm, 14-16 striae/10 mm; 16, A', cnjptocephala var. exilis, 14 X 4.5 mm, 20 striae/ 10 mm; 17, N. cnjptocephala var. veneta, 15 X 5.5 mm, 14-15 striae/ 10 mm; 18, N. cnjptoce- phala var. veneta, 24 X 7 mm, 13-14 striae/10 mm; 19, N. halophila f. tennirostris, 43 X 8.5 mm, 26 striae/10 mm; 20, A. radiosa, 60 X 11 mm, 9-12 striae/ 10 mm. All photographs are 2000X. 216 Great Basin Naturalist Vol. 42, No. 2 food webs. The annual production of the dia- toms may exceed the production of the other algae because of their faster growth rates. The higher production of diatoms is not read- ily evident because grazers often keep their biomass low (Minshall 1978). Interactions between the littoral and planktonic commimities exist, though the ex- tent of this interaction is difficult to assess. Planktonic species were found in the per- iphyton, and many littoral raphoid pennate diatoms occurred commonly in the plankton. Most freshwater phytoplankton are thought to have a neritic phase in which they dwell on the bottom, often in a resting stage (Pa- trick and Reimer 1966). This neritic phase would partly explain the occurrence of 21 22 23 phytoplankton in near shore areas, though drift and settling are also factors. Likewise, many attached algae may become unattached and drift with the plankton, which could be adaptive by helping increase their distribution. The data collected during this study in- dicate that Bottle Hollow Reservoir is a me- sotrophic to mesotrophic-eutrophic body of water. There are several evidences for this conclusion. First, biotic diversity is higher than in most eutrophic systems in the same region but lower than in many oligotrophic systems. The littoral samples with high num bers of species and absence of dominants in- dicate fairly unpolluted waters. Second, the successional pattern is not characteristic of ""^ mm^3 27 3^ 25 f * L ^ 28 29 30 31 Figs. 21-31. Diatom spp.: 21, Gornphonenia oUvaceum, 23 X 6 mm, 13-15 striae/10 mm; 22, Gomphonema parvu- him, 24 X 5.5 mm, 14-16 striae/10 mm; 23, G. intricatum, 17 X 4 mm, 12-13 striae/10 mm; 24, G. intricatum, 28 X 5 mm, 10-14 striae/10 mm; 25, G. instabilis, .34 X 7 mm, 12-18 striae/10 mm; 26, Nitzschia romana, 20 X 3.5 mm, 26 striae/10 mm, 9-10 fibulae/10 mm; 27, Amphora perpusilla, 11x3 mm, 19 striae/10 mm; 28, Cymbella microcephala, 15 X 4.5 mm, 26-27 striae/ 10 mm; 29, Nitzschia valdestriata, 10 X 2.5 mm, 10 striae/ 10 mm, 10 fibulae/ 10 mm; 30, N. sinuata var. tabellaria, 18 X 7 mm, 20 striae/10 mm, 5-6 fibulae/10 mm, 31, N. recta, 52 X 6 mm, 7-9 fibulae/10 mm. All photographs are 2000X. June 1982 JOHANSEN ET AL.: AlGAL POPULATIONS 217 eutrophic waters because blue-green algae do not play an important role. In the plankton of most eutrophic lakes and reservoirs of tem- perate regions, the large diatoms and chryso- phytes are succeeded by blue-green species in late summer, particularly Aphanizomenon flos-aquae and species of Anahaena (Wetzel 1975, Whiting et al. 1978). Such succession to Cyanophyta did not occur in Bottle Hollow Reservoir. Blue-green algae were also an in- significant part of the periphyton. Third, most diatoms we encountered (some of which are used as water quality indicators), were typical of mesotrophic waters. The two dom- inants, Asterionella formosa and Fragilaria crotonensls, are considered indicative of me- sotrophic to eutrophic water (Lowe 1974, Wetzel 1975). It should be mentioned that several species we collected often indicate eutrophic water, specifically Fragilaria vau- cheriae, Naviciiki cryptocephala var. veneta, Nitzschia minutula, Nitzschia palea, and Nitzschia paleacea (Lowe 1974). Never- theless, we have observed in our studies that these are opportunistic species that occur throughout western North America in a wide variety of habitats (Anderson and Rushforth 1976, Benson and Rushforth 1975, Johansen and Rushforth 1981, Lawson and Rushforth 1975, St. Clair and Rushforth 1976, 1978). When these species dominate a system to the exclusion of more mesotrophic organisms, they provide important evidence for eu- trophy. When they are present in lower num- bers, together with high numbers of other al- gal species (conditions we found in Bottle Hollow Reservoir) they do not necessarily in- dicate eutrophic conditions. The fourth confirmation of mesotrophic to mesotrophic-eutrophic water is the assem- blage of saprobic indicator diatoms. The saprobien spectrum was first proposed by Kolkwitz and Marsson (1908) and is a tool for assessing water quality with respect to organ- ic loading and pollution. All diatoms in Bottle Hollow Reservoir were checked against Lowe (1974). There were 23 mesosa- probic taxa, 28 oligosaprobic taxa, 12 sapro- xenous taxa, and one saprophobic species. This assemblage is evidence for water that of- ten has a moderate to high amount of dis- solved organic nutrients. It also indicates. however, that there are periods when oxida- tion is complete and water is quite "clean." Using Lowe (1974), it was also discovered that the majority of the diatoms are alkaliphilous. Our studies of Bottle Hollow Reservoir have shown that the biological water quality of this body of water is quite good, particu- larly when compared to eutrophic systems in the same region. Even so, because of the high amounts of nutrients found naturally in the rocks of the drainage basins of eastern Utah, care must be taken to limit the human-caused introduction of pollutants into this system. We believe Bottle Hollow Reservoir has the potential to maintain a healthy fishery but also has the potential for rapid deterioration toward eutrophy. Literature Cited Anderson, D. C, and S. R. Rushforth. 1976. The cryptogamic flora of desert soil crusts in southern Utah, USA. Nova Hedwigia 28:691-729. Benson, C. E., and S. R. Rushforth. 1975. The algal flora of Huntington Canyon, Utah, USA. Biblio- theca Phycologica 18:1-177. Camburn, K. E., R. L. Lowe, and D. L. Stoneburner. 1978. The haptobenthic diatom flora of Long Branch Creek, South Carolina. Nova Hedwigia 30:149-279. Foged, N. 1959. Diatoms from Afghanistan. Biol. Skr. Det Kongl. Dan. Vid. Selsk. ri(l):l-95. 1974. Freshwater diatoms in Iceland. Bibliotheca Phycologica 15:1-118. HusTEDT, F. 1930. Bacillariophyta (Diatomeae). In A. Pascher, Die Susswasser-Flora Mitteleuropas, Heft 10, Gustav Fisher, Jena, Germany. 468 pp. 1949. Susswasser-Diatomeen aus dem Albert-Na- tionalpark in Belgisch-Kongo. Exploration du Pare National Albert, Mission H. Damas (1935-19.36) 8:1-199. Inst. Pares Nat. Congo Beige, Hayez, Bruxelles. Johansen, J. R., and S. R. Rushforth. 1981. Diatoms of surface waters and soils of selected oil shale lease areas of eastern Utah, USA. Nova Hedwigia 34:333-389. Kolkwitz, R., and M. Marsson. 1908. Okologie der pflanzlichen Saprobien. Ber. Deut. Bot. Ges. 26:.505-519. Lawson, L. L., and S. R. Rushforth. 1975. The diatom flora of the Provo River, Utah, USA. Bibliotheca Phycologica 17:1-149. Lowe, R. L. 1974. Environmental requirements and pol- lution tolerance of freshwater diatoms. Environ- mental Monitoring Series, U.S. Environmental Protection Agency, Report No. EPA-670/4-74- 005. .334 pp. Merritt, L., S. Rushforth, J. Johansen, R. Heck.man, R. Winget, and W. Brimhall. 1980. Assessment 218 Great Basin Naturalist Vol. 42, No. 2 of water quality in Bottle Hollow Reservoir. Re- port to Ute Indian Tribe, Duchesne Co., Ut. 180 pp. MiNSHALL, G. W. 1978. Autotrophv in stream ecosys- tems. BioScience 28(1 2): 767-771. Patrick, R., and C. W. Reimer. 1966. The diatoms of the United States, I. Acad. Nat. Sci. Philad. Mon 13. 1-688 pp. Pearsall, W. H. 1932. Phytoplankton in the English Lakes. II. The composition of the phytoplankton in relation to dissolved substances. I. Ecology 20(2):241-262. Porter, K. G. 1977. The plant-animal interface in fresh- water ecosystems. Am. Scientist 65:159-170. St. Clair, L. L., and S. R. Rushforth. 1976. The dia- toms of Timpanogos Cave National Monument, Utah. Amer. J. Bot. 6.3(l):49-59. 1978. The diatom flora of the Goshen Playa and wet meadow. Nova Hedwigia 29:191-229. Wetzel, R. G. 1975. Limnology. W. B. Saunders Com- pany, Philadelphia, Penn. 743 pp. Whiting, M. C, J. D. Brotherson, and S. R. Rushforth. 1978. Environmental interaction in summer algal communities of Utah Lake. Great Basin Nat. 38(1):31-41. WiMPENNY, R. S. 1973. The size of diatoms. V. The ef- fect of animal grazing. J. Mar. Bio. Assoc. United Kingdom 53:957-974. HERPETOLOGICAL NOTES FROM THE NEVADA TEST SITE Wilmer W. Tanner' Abstract.- During the years 1965-1971, considerable data were gathered that included information concerning species not previously reported. These included Chionactis occipatalis talpina, Coleonyx variegatus utahensis, Crota- phytus coUaris bicinctores, Cnemidophorus tigris tigris, and Sauromalus obesus obesus. Although complete informa- tion concerning their life histories is not reported, some information concerning growth and reproduction is included. The following notes were made by Mr. Ronald L. Morris, Dr. John E. Krogh, or the author and represent our findings concerning species found in our NTS study plots not re- ported previously. Several of our study plots provided some data on species that were not intensively studied or the limited data did not seemingly justify, at that time, consideration. It was our intent to gather additional data; however, this was not possible, and I am, therefore, presenting those data which are considered to have value. Study plots were originally established for the express purpose of examining in as much detail as possible the life histories and habi- tats of the more abundant species. Thus, Rainier Mesa provided data on Sceloporus oc- cidentalis and Uta stansburiana (Tanner and Hopkin 1972, Tanner 1972). Three study plots in Frenchman Flat examined Crota- phytus wislizeni (Tarmer and Krogh 1974), Phrynos&ma platyrhinos (Tanner and Krogh 1973), and Callisaurus draconoides (Tanner and Krogh 1975). Data from the Mercury Valley plot were included in some of the re- ports listed above. In three of the five study plots, can pitfall traps were used; this enabled us to at least sample most of the species, particularly small lizards and snakes, that occurred in the habi- tat being studied. In the other two plots liz- ards were caught and marked by means of a noose. At the Frenchman Flat Plot 2 (a rocky hill completely surrounded by desert flats and sit- uated west of the Mercury highway and south of the Kane Springs road), small popu- lations of Crotaphytus collaris and Sauro- radiis obesus were studied. At the Mercury Valley plot some data were obtained for Coleonyx variegatus and Chionactis occipa- talis. Although only fragmentary data are presented, it does seemingly have merit. Chionactis occipatalis talpina Klauber We marked 62 individuals; of these 5 were recaptured once, and one twice. All recap- tures were within 40 to 180 feet of the origi- nal capture, and 4 were recaptured two years after the original capture. Although these data do not substantiate a home range, they do indicate a relatively small "homing" area for this species. The can traps were set 40 feet apart in rows of 10 traps, and in rows numbered from A to T. Number 16, a young adult (S-V 225 mm), moved from G-8 (15 June 1966) to F-3 (5 August 1967) and to C-5 (5 June 1968). The smallest individual marked was a hatchling marked 28 August 1969 with a S-V of 109 mm. The largest female was 291 mm S-V and the largest male was 287 mm. Indi- viduals were considered adults if they were 250 or more in S-V length. Weight of adults varied with size and, in females, before eggs were layed. In adults, weights ranged from 6 to 10.4 grams. Growth Three of those recaptured were juveniles or subadults and show the following growth: 'Life Science Museum, Brigham Young University, Provo, Utah 84602. 219 220 Great Basin Naturalist Vol. 42, No. 2 No. 7 marked 15 August 1965 S-V 215 mm. Recaptured 5 June 1967 S-V 236 mm growth, 21 mm in 2IV2 months. No. 12 marked 3 June 1966 S-V 239 mm. Recaptured 5 June 1968 S-V 251 mm growth, 12 mm in 24 months. No. 16 marked 15 June 1966 S-V 225 mm. Recaptured 5 August 1967 S-V 250 mm. Re- captured 5 June 1968 S-V 251 mm growth, 25 mm in 13'/3 months. Growth after individuals attain 250 mm in S-V length is seemingly slowed to a few mm per season, or, as in number 16, growth of no more than 1 mm in 24 months. Our data in- dicate that subadults may grow 0.5 to 2 mm per month, and show that from hatching to the largest adults they provide a S-V growth of approximately 180 mm. We do no t have any data on longevity. Coleonyx variegatus utahensis Klauber At the Mercury Valley study plot we marked 115 individuals, and of these 33 were recaptured; 23 once, 7 twice, 2 three, and one five times. These data do not provide suf- ficient information to establish with certainty a home range size for this species. We con- clude this in spite of the fact that many indi- viduals were recaptured not far from their original capture, but only a few were recap- tured more than twice. Those recaptured three or more times during two or more years do indicate a relatively small home range. Females were gravid during July and early August in 1965. Hatchlings were seen in Au- gust. The earliest seen by us was 10 August 1966. Hatchlings during August (10th to 30th) from 1965-1969 range in size from 32 to 40 mm. It is assumed that the larger hatch- lings seen in late August had been extant for several weeks, thus accounting for their larger size. Twelve juveniles caught in June (5th to 29th) ranged in size from 45 to 58 mm. Growth of hatchlings continues into September and begins again in April or early May; this accounts for the 10 to 15 mm of added growth seen in June. Growth continues so that by August most juveniles have reached a S-V length of at least 60 mm, and by the next June are be- tween 65 to 70 mm, most nearing 70 mm. The largest female individual seen measured 73 mm in S-V, was gravid, and weighed 6 grams on 1 August 1965. Crotaphytus coUaris bicinctores Smith & Tanner Great Basin Collared Lizard Five adults were marked in 1965-1966 at the Frenchman Flat Plot 2. Each of these were recaptured or observed many times un- til the plot was closed in 1971. The habitat is a rocky ridge approximately 500 yards long and about half as wide. It rises to a peak in the middle and was used during the open air atomic testings as an observation point. On its top was a pole, which we refer to as the flag pole. Because of the openness of the hab- itat, the lizards moved over large areas with- in a loosely considered home range; this was quite in contrast to the limited space avail- able to hzards in the study by Fitch (1956). Number 5 was observed during five years and traversed the southeastern side of the ridge, a linear distance of 680 feet. Although the others were not observed to travel this far, the home ranges, if indeed such occur, were large. There was an overlapping of ranges, even by males, a condition perhaps related to the openness of the large range and the small number of individuals present. We assumed the small population to be related to the food supply. We did not see many insects, and a population of Sceloporus majester shared the habitat and the food. The lizards were easily spotted on rocks, usually on those higher than others nearby. They were not easily frightened, so we could observe them at close range. On 9 June 1966 we observed number 2, an adult male, vigor- ously active around a large rock near the flag pole (F-10). As we approached the rock, we noted a swarm of flying ants around the rock, and as the ants fluttered and tumbled over and down the rock, they were eaten by the lizard. At times he would feed on four to six ants in rapid succession, when they were available at the base of the rock. He would jump to capture flying ants if others were not available on the ground. He was relatively tame and paid little attention to us, even though we were only a few feet away. June 1982 Tanner: Herpetological Notes 221 On 1 July 1966, number 1, an adult female, was recaptured at G-6; on being released she moved up the ridge to F-10 and soon there- after was observed eating flying ants that were still swarming on a rock near the flag pole. The basking lizards were seen to leave their rock perch and chase a short distance, and then return. Number 5, an adult female, was observed on a rock near H-11 (about 100 feet from the flag pole); after observing for some time, we threw small rocks near her, and each time she responded by rushing to the spot where the rock lit. We were about 40 feet away, and by continuing to throw pebbles we were able to draw her right up to us. Each time after chasing a rock she would perch on a nearby rock and watch. Our ob- servations of feeding activities compare sim- ilarly to those of Fitch (1956), and indicate a series of similar activity and feeding behaviors. We did not see hatchings at this plot, al- though each of the females was observed to be gravid on 29 June 1966. Growth was ob- served in three, with numbers 2 and 3 (who were 103 and 89 mm in S-V length) showing no growth. Number 5 was 79 mm in S-V on 12 June 19667, and had grown to 95 mm on 15 May 1971. Weight during this same peri- od increased from 21 to 28.8 grams; only gravid females weighed more (number 3 on 12 June 1966 weighed 33 grams). Although we do not have data for com- plete growth and longevity, we do recognize this species as one with perhaps as long or longer life span than other lizards at this plot, except for Sauromalus. By comparing the size of number 5 (79 mm) to specimens in our preserved collection, we were not able to de- termine if she was nearing one or two years. Her size would represent only one year if based on data for the eastern subspecies (Fitch 1956). The study by Fitch shows rapid growth, with some individuals reaching full adult size in one year, an indication of abun- dant food and perhaps more favorable cli- matic conditions. The desert foothills and valleys of the Frenchman Flat area (Tanner and Hopkin 1972, Fig. 4) are dry and hot for most of June through September, with only an occasional thimderstorm. This not only af- fects the activities of the lizards, but also seemingly dries up the vegetation and may reduce the availability of insect food. In spite of the fact that the two studies are of two widely separated, distinct subspecies, we be- lieve that environmental factors are extreme- ly important in providing the differences in growth rate. When last seen, number 5 (15 May 1971) was in apparent good health and was either in her sixth or seventh year. Cnernidophonis tigris tigris Baird & Girard Great Basin Whiptail Our field data concerning reproduction, growth, and longevity confirm previous stud- ies such as those of Tanner and Jorgenson (1963), McCoy and Hoddenbach (1966), Burkholder and Walter (1973), and Turner, Medica, Lannom, and Hoddenbach (1969). The study by Tanner and Jorgenson (1963) suggested that aestivation, or early hiberna- tion, occurs in early summer. We have ob- served this during a five-year study of popu- lations at our study plots in Frenchman Flat and Mercury Valley. At both plots there was a noticeable reduction in the number of adults seen beginning in mid-July; the last adults were seen occasionally until mid- August. The latest record we have of an adult (92 mm S-V) was 26 August 1965. During August, hatchlings and second-year juveniles are seen, but by September only the year's hatchlings, ranging in size from 40 to 55 mm in S-V length, are seen. This does not mean that an occasional adult may not be seen, but our record indicates that a decided reduction in activity occurs each year beginning in mid-July. By far the greatest activity is in May and June. We also noted that adult individuals marked in June or July, and then recaptured in April or May (soon after becoming active the next year), invariably had lost several grams of weight. Examples are toe-mark number 2-4, 8 June 1967, S-V 91, weight 24.8 grams; recaptured 7 April 1968, S-V 88, weight 20.8 grams. This individual was seen again on 4 May 1968 with a S-V of 93 and 23.1 grams. Number 1-6 marked 7 June 1966, S-V 99, weight 26 grams, recaptured 7 April 1968, S-V 96, weight 25 grams. Our data sug- gest that the early hibernation has the effect 222 Great Basin Naturalist Vol. 42, No. 2 of reducing size and weight, which is rapidly regained by adults in June. Sauromalus obesus obesus Baird Western Chuckwalla A population of chuckwallas were seen in Mercury Pass as we traveled from Mercury to our study plots in Frenchman Flat and Rainier Mesa. As time and opportunity was available, we marked 45 and recaptured 17; 9 once, 5 three, and 3 four times. No attempt was made to determine home range size, al- though our records indicate that certain rocks with suitable cracks and holes served as a home base around which they foraged and sunned. As we continued to mark and recapture, we noted considerable growth in both size (mm) and weight (grams). Three males pro- vided the following data: Number 2 on 30 June 1965 had S-V 175 mm and weighed 238 grams. On 29 June 1966 S-V 250 and weighed 315 grams. On 4 May 1968 only 2 mm of additional growth had occurred, but he had gained 80 additional grams to weigh 395 grams. Number 4 on 16 July 1965 had a S-V of 150 and weighed 159 grams. On 28 May 1966 the S-V was 173 and weighed 210 grams. On 11 June 1968 he was S-V 185 and weighed 287 grams. Number 6 on 21 July 1965 was S-V 154 and weighed 146 grams. On 28 June 1967 number 6 (two years' growth) was S-V 181 and weighed 251. This individual had aver- aged over 50 grams per year, but only 27 mm in S-V length. We noted that the greatest growth of those marked occurred in the first tliree years; this is particularly true for the S-V length. After three years, most growth was in terms of added weight. Males were larger than fe- males. The largest male (number 7) seen weighed 432 grams and was at least five years old. This individual gained only 39 grams in 3 years (393 grams on 28 May 1966 to 432 grams on 13 June 1968). Our data in- dicate that, after the S-V length of 200 mm is reached, a considerable slowing in length growth occurs; but, as noted above, increase in weight continues. We noted courtships in May and early June, and that females were gravid in July. Three clutches of eggs were laid in July, and ranged in number from 4 to 14. Literature Cited BuRKHOLDER, G. L., AND J. M. Walker. 1973. Habitat and reproduction of the desert whiptail lizard, Cnemidophorus tigris Baird and Girard at the northern part of its range. Herpetologica 29:76-83. Fitch, H. S. 1956. An ecological study of the collared lizard {Crotaphytus collaris). Univ. of Kansas Publ. Mus. Nat. Hist. 8(3):213-274. McCoy, C. J., and G. A. Hoddenbach. 1966. Geo- graphic variation in ovarian cycles and clutch size in Cnemidophorus tigris (Teiidae). Science 154:1671-1672. Tanner, W. W. 1972. Notes on the life history of Uta s. stansburiana. BYU Sci. Bull, Biol. Ser. 15(4):31-39. Tanner, W. W., and J. M. Hopkin. 1972. Ecology of Sceloporus occidentalis longipes Baird on Rainier Mesa, Nevada Test Site, Nye County, Nevada. BYU Sci. Bull., Biol. Ser. 15(4): 1-31. Tanner, W. W., and C. J. Jorgenson. 1963. The appli- cation of the density probability function to de- termine the home ranges of Uta stansburiana stansburiana and Cnemidophorus tigris tigris. Herpetologica 19(2): 105-1 15. Tanner, W. W., and J. E. Krogh. 1973. Ecology and natural history of Phrynosoma platyrhinos, Ne- vada Test Site, Nye County, Nevada. Herpeto- logica 29(4):327-342. 1974. Ecology of the leopard lizard, Crotaphytus wislizeni at the Nevada Test Site, Nye County, Nevada. Herpetologica 30(l):63-72. 1975. Ecology of the zebra-tailed lizard Calli- saurus draconoides at the Nevada Test Site. Her- petologica 31(3):302-316. Turner, F. B., P. A. Medica, J. R. Lannom, and G. A. Hoddenbach. 1969. A demographic analysis of fenced populations of the whiptail lizard Cne- midophorus tigris, in southern Nevada. South- west. Natur. 14:189-201. NEW SPECIES OF AMERICAN BARK BEETLES (COLEOPTERA: SCOLYTIDAE) Stephen L. Wood' ,\bstract.— The following species of Scolytidae (Coleoptera) are described as new to science: Amphicranus splen- dens, Araptus tnicwpilosus (Mexico), Araptus morigerus (Panama), Araptus placetiilus (Mexico), Chramesiis bispinus (Colombia), Cnesinus aquihuai (Mexico), Cnesinus atrocis (Panama), Cnesinus meris (Colombia), Dendrocranulus gracilis, Hi/lustes retifer, Hylocurus prolatus, Micracis burgosi, Phloeotribus perniciosus, Pseudothysanoes fimbriatus, Pscudothi/sanoes pini, Scolytodes pilifer, Scohjtus binodus (Mexico). While my taxonomic monograph of the Scolytidae of North and Central American (Wood 1982) was in press, several species new to science came to my attention. The following pages report 17 of these species in the genera Amphicranus (1), Araptus (3), Chramesus (1), Cnesinus (3), Dendrocranulus (1), Hylastes (1), Hylocurus (1), Micracis (1), Phloeotribus (1), Pseudothysanoes (2), Scoly- todes (1), and Scolytus (1) from Mexico (13), Panama (2), and Colombia (2). Most of the Mexican material was received through Dr. T. H. Atkinson from the Colegio de Post- graduados, Institucion de Ensenanza y In- vestigaciones Agricolas, Chapingo, Mexico. Amphicranus splendens, n. sp. This species belongs to the melanura spe- cies group of the genus and is the most spe- cialized known species in that group. It is distinguished from argutus Wood by the larger size, by the reticulate, dull frons, by the modified anterior margin of the pro- notum, by the very different elytral declivity, and by other characters. Male.— Length 3.1 mm (paratypes: male 3.5 mm, female 3.0-3.2 mm), 3.2 times as long as wide; color very dark brown. Frons broadly, uniformly convex; surface dull, very finely reticulate-granulate, punc- tures and vestiture virtually obsolete; a few setae at margins of eyes, epistomal margin ornamented by moderately abundant, long hair. Antennal club 1.7 times as long as wide, 1.7 times as long as scape; sutures 1 and 2 clearly indicated, slightly procurved; club densely covered by minute hair. Pronotum 1.4 times as long as wide; sides straight and parallel on basal half, gradually tapering to angle on anterior margin; ante- rior margin straight on median half, formed by an elevated, continuous costa, meeting lat- eral margin at an abrupt, obtuse angle; sum- mit indefinite, anterior to middle; low, al- most scalelike asperities restricted to slightly less than anterior half; surface reticulate ex- cept for shining asperities and transverse rugae on more than posterior third; punc- tures minute, moderately close. Glabrous. Elytra 1.8 times as long as wide, 1.3 times as long as pronotum; sides straight and paral- lel to base of declivital processes, posterior margin almost straight, feebly incised; disc shining, almost smooth, with rather numerous very feebly impressed lines, punctures min- ute, confused, rather close. Declivity sub- vertical; ventrolateral margin on more than lower third of a complete circle acutely, moderately explanate, entire at suture; upper margin not elevated at suture, interstriae 2 armed by a small, pointed tubercle borne on an obtuse elevation, summit of this elevation continuing to major process; major process occupying area of about interstriae 4 to 5, its lower margin at middle of declivity, its later- al surface continuing contour and sculpture of disc, its apex bluntly rounded, its free pro- jection equal in length to width of antennal club, its mesal surface armed by a coarse, subapical denticle; face of declivity strongly, broadly concave, punctures fine, confused, 'Life Science Museum and Department of Zoology, Brigham Young University, Provo, Utah 84602. 223 224 Great Basin Naturalist Vol. 42, No. 2 obscure. Vestiture fine, short, obscure on de- clivital face, a few longer setae on lateral areas. Female.— Similar to male except upper third of elytral declivity occupied by large, rounded protuberance in place of major pro- cess, small denticle at base on interstriae 2 present; antennal club without long setae on posterior face. Type material.— The male holotype, fe- male allotype, and three paratypes were taken at San Rafael, Mexico, Mexico, 4-IX- 1981, 2400 m, S-240, Quercus, T. H. Atkin- son and A. Aquihua. The holotype, allotype, and paratypes are in my collection. Araptus micropilosus, n. sp. This species keys to tenuis (Blackman) in Wood (1982:933). It is distinguished by the very different female frons and by the strong- ly reticulate pronotum. Female.— Length 1.4 mm (paratypes 1.2-1.5 mm), 2.5 times as long as wide; color dark reddish brown. Front flattened on a subcircular area oc- cupying more than median three-fourths of area between eyes from epistoma to vertex, central half of flattened area micropunctate and ornamented by dense, minute micropile, surrounding area more coarsely punctured and bearing a rather sparse brush of long hair. Antennal club subcircular, without defi- nite sutures (suture 1 apparently marked on some specimens). Pronotum 1.05 times as long as wide; widest on basal fourth, tapered anteriorly, then rather narrowly rounded on finely ser- rate anterior margin. Summit indefinite, be- hind middle; posterior areas strongly reti- culate, broad median line impunctate; remainder of disc with sparse minute punc- tures, these largely replaced by fine granules. Largely glabrous. Elytra 1.6 times as long as wide; sides al- most straight and parallel on basal two-thirds, rather broadly rounded behind; striae not im- pressed except 1 slightly, punctures small, shallow, in rows; interstriae about twice as wide as striae, smooth and shining except for rather abundant irregular lines, pmictures ab- sent. Declivity rather steep, convex; suture weakly elevated, striae 1 narrowly, rather deeply impressed, 2 as high as 1, and 1, 2, and 3 each with a row of very minute gran- ules. Vestiture largely abraded on type, con- sisting of rows of erect, stout, blunt, inter- strial bristles on declivity, each bristle about as long as distance between rows, spaced within a row by length of a bristle. Male.— Similar to female except frons convex coarsely, closely punctured, a feeble median carina on middle half, vestiture fine, sparse, inconspicuous. Type material.— The female holotype, male allotype, and seven paratypes were taken at Rancho Tepetates, Km 35 on the Veracruz— Xalapa highway, Veracruz, Mexi- co, 12-VI-1979, T. H. Atkinson. The holo- type, allotype, and paratypes are in my collection. Araptus blanditus Wood Araptus blanditus Wood, 1974:47 (Holotype, female; Fortin de las Flores, Veracruz, Mexico; Wood Coll.) In my collection three species have been confused under this name. The unique female type has the frons moderately concave, the pronotal disc is rather long, smooth, brightly shining, and without impressed points but with rather large, almost round, rather wide- ly spaced punctures, the discal strial punc- tures are almost all in rows, declivital inter- striae 2 is not wider than 1 and ascends rather conspicuously laterally, with the pimc- tures of striae 2 comparatively coarse, and the antennal club is comparatively small and slender (1.5 times as long as wide). As point- ed out below, slight but consistent differences separate this species from the two that follow. Araptus morigerus, n. sp. This species was incorrectly reported (Wood 1982:951) as blanditus Wood. It is distinguished from blanditus by the charac- ters described below. Female.— Length 1.9 mm (paratypes 1.7-2.0 mm), 2.6 times as long as wide; color dark reddish brown. Frons as in blanditus, except very slightly more strongly impressed, a small median tu- bercle on epistoma (obscure in a few speci- mens). Antennal club as in blanditus. June 1982 Wood: New American Scolytidae 225 Pronotiim as in hlanditus except with mod- erately abundant impressed points, punctures smaller, closer, mostly elongate. Elytra as in hlanditus except strial punc- tures on anterior half of disc moderately con- fused, surface with moderately abundant im- pressed points; declivity shorter, steeper, punctures on striae 2 smaller, interstriae 2 wider, more nearly flattened; setae much finer. Male.— Similar to female except frons broadly convex, a feeble transverse impres- sion just below middle, without any other im- pressions, punctures fine, rather abundant, vestiture fine, short, uniformly distributed; epistoma straight. Type material.— The female holotype, male allotype, and 33 paratypes were taken near Cerro Punta (labeled Volcan Chiriqui), Panama, 11-1-1964, 5500 ft. No. 376, broken branch, by me. The holotype, allotype, and paratypes are in my collection. Araptus placetulus, n. sp. This species is distinguished from hlanditus Wood by the characters described below. Female.— Length 1.7 mm (males 1.9 mm), 2.4 times as long as wide; color very dark brown. Frons similar to hlanditus except less strongly impressed, surface more nearly gran- ular, vestiture shorter (about two-thirds as long). Antennal club larger, stouter (1.3 times as long as wide). Pronotuin as in hlanditus except anterior slope more gradual, disc slightly shorter, with numerous impressed points, punctures slightly smaller, much closer, elongate. Elytra as in placetus on disc; declivity about as in hlanditus except interstriae 2 more strongly impressed, almost flat, as wide as 1; vestiture fine as in morigerus. Male.— Similar to female except frons convex, a slight median callus at upper level of eyes, a transverse impression just below middle; epistoma slightly recurved, median third weakly elevated, slightly impressed in lateral areas before bases of mandibles, punc- tures very fine in central area on lower half, longer laterally and above. Type material.— The female holotype, male allotype, and one male paratype were taken at Uruapan, Michoacan, Mexico, 1-XI- 1980, 1600 m, S-149, Aguacate, T. H. Atkin- son and A. Equihua. The holotype, allotype, and paratype are in my collection. Chramesus hispinus, n. sp. This species is unique in the genus. It is distinguished by the bicolored pattern of scales and by the pair of large, hornlike spines on the elytral declivity. Male.— Length 1.8 mm (paratypes 1.9-2.2 mm), 1.9 times as long as wide; color a some- what variable pattern of pale and dark scales. Frons with a low, transverse elevation at level of antennal insertion, shallowly concave below this elevation, moderately concave above to upper level of eyes; surface sub- reticulate and finely punctured; vestiture of short stout setae, erect and slightly longer on lateral margins of upper area, epistoma with a brush of longer, yellow setae. Antennal scape with a tuft of long, yellow hair; club comparatively small for this genus, an appar- ent obscure surface indication of suture 1 present on anterior face. Pronotum 0.77 times as long as wide; widest at base, sides moderately arcuate to rather broadly rounded anterior margin, a slight constriction just before anterior mar- gin; surface smooth, shining, punctures small, close. Vestiture of short, abundant scales, those on basal third and lateral margins pale, dark in central and anterior areas; a few pale scales scattered in dark areas. Elytra 1.3 times as long as wide, 1.8 times as long as pronotum; sides almost straight and parallel on basal two-thirds, rather broadly rounded behind; striae not impressed, punc- tures small, distinctly impressed; interstriae smooth, shining, about four times as wide as striae, lateral half of 1 to mesal half of 2 shal- lowly sulcate from base, broadening and deepening to become impression of declivity. Declivity commencing slightly behind middle, moderately steep; interstriae 1 dis- tinctly elevated, strongly, rather broadly im- pressed from 1 to 3 on upper half, shallowly, more broadly impressed below; interstriae 3 armed slightly above middle by a pair of very coarse, hornlike spines directed caudomesad and slightly dorsad, each equal in length to 226 Great Basin Naturalist Vol. 42, No. 2 width of antennal club. Vestiture of inter- strial cover of short ground scales, each slightly longer than wide, and rows of erect bristles, those near base only slightly longer than ground scales, becoming longer and more slender posteriorly except on central part of lower declivity, these setae continue to apex of spines. Female.— Similar to male except frons less strongly impressed; tuft of hair on scape con- spicuously smaller. Type material.— The male holotype, fe- male allotype, and five paratypes were taken at Tenerife, Valle, Colombia, 1-81 (three par- atypes IX-80), tallos de curuba (passion flow- er vine stems, Passiflora mollisima), Patricia Chacon. Cnesinus aquihuai, n. sp. This species is distinguished from atavus Wood by the slightly larger, stouter body, and by the very different frons that is de- scribed below. Female.— Length 2.5 mm, 2.2 times as long as wide; color dark reddish brown. Frons broadly, subconcavely impressed be- tween lateral margins from epistoma to up- per level of eyes; upper margin of impressed area at upper level of eyes abrupt, sub- carinate; floor of impressed area reticulate, ornamented on a triangular area by special, compressed, reddish brown setae, base of triangle on epistoma, occupying median two- thirds, its apex on median line two-thirds dis- tance toward upper level of eyes. Pronotum and elytra essentially as in atavus. Male.— Similar to female except frontal impression somewhat irregular, ending grad- ually well below upper level of eyes; special ornamental setae absent, those present in lat- eral areas above epistomal area longer, yellowish. Type material.— The female holotype, male allotype, and two female paratypes were taken between Cuetzalan and Pasa del Jardin, Puebla, Mexico, 5-V-1981, 550 m, S- 224, by T. H. Atkinson and A. Aquihua. The holotype, allotype, and paratypes are in my collection. Cnesinus atrocis, n. sp. This species is distinguished from the allied bicolor Eggers by the very different female epistomal callus and its setal ornamentation, by the more deeply impressed striae, by the larger strial punctures, and by the near ab- sence of interstrial punctures except for those bearing the uniseriate rows of erect bristles. Female.— Length 2.3 mm, 2.7 times as long as wide; color reddish brown. Frons as in bicolor except for epistomal callus; epistomal callus small, its upper mar- gin forming a rather high, subacute, almost straight, transverse carina on more than me- dian two-thirds, upper slope of callus (dorsad of crest of carina) ornamented by a single row of reddish, compressed specialized setae extending without interruption full length of carina. Pronotum as in bicolor except punctures somewhat larger and more strigose, with lim- ited confluence of strigosities on middle third. Elytra about as in bicolor except striae more deeply, abruptly, narrowly impressed; bristle-bearing interstrial punctures almost uniseriate and distinctly crenulate, supple- mental punctures obsolete, interstrial bristles coarser, rarely abraded. Type material.— The female holotype was taken near Cerro Punta (labeled Volcan Chiriqui), Panama, 11-1-1964, 5500 ft No. 388, from the twig of an unidentified sapling, by me. The holotype is in my collection. Cnesinus meris, n. sp. This species is distinguished from bisul- catus Schedl by differences in the epistomal callus and its setal ornamentation and in the sculpture of the pronotum. Female.— Length 2.4 mm, 2.6 times as long as wide; color dark reddish brown. Frons as in bisulcatus except epistomal cal- lus more abruptly elevated, its surface smooth, dull on large triangular area, upper margins of flattened area bearing one row of short, compressed, reddish brown ornamental bristles (in bisulcatus callus smaller, its mar- gins less abrupt, upper two-thirds covered by ornamental setae). June 1982 Wood: New American Scolytidae 227 Pronotum as in bisulcatus except punc- tures averaging smaller, not as close (very similar). Male.— Similar to female except frontal callus greatly reduced, its ornamental setae finer, yellowish, more nearly normal. Type material.- The female holotype, male allotype, and 12 paratypes were taken at La Cumbre, Valle, Colombia, 3- VI- 1959, en cafe, by N. Muiioz. The holotype, allo- type, and paratypes are in my collection. Dendrocranulus gracilis, n. sp. This species is distinguished from the allied macilentus (Blandford) by the much smaller size, by the larger pronotal and elytral punc- tures, iDy the very different female frons, and by other characters described below. Female.— Length 1.4 mm (paratypes 1.3-1.6 mm), 3.2 times as long as wide; color black. Frons broadly convex except flat on triangular area occupying median half at epistoma to obtuse apex well above upper level of eyes, this shining area minutely, rather densely punctured and ornamented by very fine, abundant, short hair, remaining area more coarsely punctured; vestiture less abundant. Pronotum 1.2 times as long as wide; sub- quadrate, all margins moderately, about equally arcuate; surface obscurely sub- reticulate; indefinite summit near middle; discal area coarsely punctured, punctures largely replaced by slender elongate calluses in lateral areas; vestiture of sparse hair. Elytra 2.0 times as long as wide, 1.7 times as long as pronotum; sides straight and paral- lel on basal three-fourths, very broadly rounded behind; striae not impressed, punc- tures rather coarse, in rows; interstriae as wide as striae, smooth, punctures uniseriate, half as large as those of striae. Declivity very steep, broadly convex, almost flat; surface sculpture as on disc; suture distinctly ele- vated, flat from striae 1 to 4, lower margin subacute to interstriae 4. Vestiture very fine, of rather short strial and conspicuous longer interstrial hair of moderate abundance. Male.— Similar to female except frons more strongly convex above, a weak trans- verse impression at level of antennal insertion, coarsely punctured and sparsely pubescent over entire surface; declivital in- terstriae 2 rather broadly, moderately im- pressed, smooth, shining, impunctate. Type material.— The female holotype, male allotype, and four paratypes were taken at Km 58 on the Xochimilco-Oaxtepec high- way, Morelia, Mexico, 22-1-1980, 1970 m, S- 19, Cucurbitaceae, T. H. Atkinson; six para- types are from Uruapan, Michoacan, Mexico, 16-V-1981, Sechium edulis, A. Equihua. The holotype, allotype, and paratypes are in my collection. Hylastes retifer, n. sp. This species keys to macer LeConte in my monograph (Wood 1982:95), although its true affinities appear much closer to mexi- canus Wood. It differs from all American species except macer by the uniformly re- ticulate pronotum and elytra. From macer it is distinguished by the stouter body, by the larger, stouter pronotum, with punctures much smaller and more abundant, and by the smaller, less strongly impressed strial punc- tures on the disc. Male.— Length 5.4 mm, 2.8 times as long as wide; color black. Frons essentially as in macer except more finely, closely punctured. Pronotum 1.1 times as long as wide; sides parallel on basal half; surface uniformly, very finely reticulate (visible at minimum of 40X), punctures small, moderately deep, abundant, spaced by distances about equal to diameter of a puncture. Elytra 1.9 times as long as wide; striae rather weakly impressed except near declivi- ty, pimctures rather small, moderately im- pressed; entire surface uniformly, finely reti- culate; interstriae distinctly wider than striae, punctures fine, confused, rather abundant, 2 on posterior half of disc wider than 1 or 3 and weakly elevated; declivity as in mexi- canus except reticulate. Type material.- The male holotype was taken at Km 54 Carretera Toluca-Morelia, Est. de Mexico, 30-X-1980, 2520 m, Pinus montezumae, T. H. Atkinson and A. Equihua. The holotype is in my collection. 228 Great Basin Naturalist Vol. 42, No. 2 Hylocurus prolatus, n. sp. This species is distinguished from long- ipennis Wood by the much weaker male frontal carina, by the less protuberant, more pubescent female frons, and by the sculpture at the basal margin of the declivity. Male.— Length 2.8 mm (paratypes 2.5-2.9 mm), 3.3 times as long as wide; color black. Frons convex, a slight transverse pro- tuberance on middle third, a weak transverse carina indicated about two-thirds distance from epistomal margin to upper level of eyes. Pronotum and elytra as in longipennis ex- cept for basal area of elytral declivity; inter- striae on posterior fourth of disc moderately convex, their summits interrupted at punc- tures (not clearly nodulate as in longipennis), their crests very briefly, longitudinally cari- nate at subabrupt margin of declivity (much more so than in longipennis; declivital inter- striae 9 more strongly elevated than in long- ipennis, its crest much less strongly tuberculate. Female.— Similar to male except frons re- sembling female longipennis, with pro- tuberance almost obsolete, median half of lower third impunctate, almost smooth, transverse area at upper level of eyes more finely punctured and ornamented by more abundant, fine, short, somewhat reddish setae; interstriae at base of declivity without longitudinal short carinae, margin less abRipt. Type material.— The male holotype, fe- male allotype, and 18 paratypes were taken at Zacapoaxia, Puebla, Mexico, 6- V- 1981, 2150 m, S-230, Canja, T. H. Atkinson and A. Equihua. The holotype, allotype, and para- types are in my collection. Micracis burgosi, n. sp. The antennal scape of the female is the most remarkable found in any scolytid. Al- though the antennal club clearly places this species in Micracis, the protibia is more slen- der than seen elsewhere in this genus and the posterior face bears a few minute tubercles as in Hylocurus. Rather than suggest synonymy of these genera on the basis of this inter- mediate species, it is arbitrarily assigned to Micracis because of the antennal structure. It shares with dimorphus (Schedl) an identical protibia and antennal club, and a secondary shaft on the female scape, but it differs in nu- merous characters, some of which are de- scribed below. Female.— Length 2.6 mm (paratypes: male 2.2-2.4 mm, female 2.3-2.8 mm), 3.3 times as long as wide; color dark brown. Frons feebly, transversely impressed at lev- el of antennal insertion, flattened below, up- per area to well above eyes weakly convex, central half smooth, brightly shining, im- punctate; lateral and upper margins finely punctured and ornamented by moderately long, rather abundant setae. Antennal scape to insertion of funicle club shaped, its length equal to about one and one-half times width of eye, its dorsal margin ornamented by a fringe of rather abundant, long setae, most of these more than twice as long as this portion of scape, scape extended on dorsoapical angle above insertion of funicle into a long, slender shaft equal in length to combined length of funicle and club (or almost twice length of basal portion of scape), basal two-thirds of this shaft as wide as funicle, apical third twice this width, shaft glabrous except dorsal margin of its apical third ornamented by a tuft of very long setae, some of these setae longer than entire antenna; funicle and club as in dimorphus, except club very slightly more slender. Pronotum 1.3 times as long as wide; out- line as in dimorphus; posterior areas sub- reticulate, shining, crenulations continuing to base, except decreasing in size on posterior third and usually with rudimentary puncture on their posterior margins in this area. Elytra 1.9 times as long as wide; sides straight and parallel on basal three-fourths, obtusely pointed behind; striae not impressed except 1 feebly, punctures moderately coarse, rather deep; interstriae one and one-half times as wide as striae, smooth, shining, punctures almost as large as those of striae, shallow, rather widely spaced. Declivity steep, convex; sculpture about as on disc ex- cept odd-numbered interstriae each with a row of about four fine tubercles, 9 moder- ately, subacutely elevated near its apex, crest of this elevation joining costal margin and continuing at descending height to sutural apex. Vestiture consisting of minute strial June 1982 Wood: New American Scolytidae 229 hair and rows of erect, coarse interstrial setae, those on disc about as long as distance between rows, almost twice as long on declivity. Protibia about as in dimorphus except tu- bercles on posterior face smaller. Male.— Similar to female except frons more strongly convex, coarsely punctured on upper two-thirds, vestiture simple, rather short, uniformly distributed; scape simple, without ornamentation. Type material.— The female holotype, male allotype, and 25 paratypes were taken at Cuemavaca, Morelos, Mexico, 2-II-1981, Delonix regia bole, A. Burgos. Phloeotribus perniciosus n. sp. This species is distinguished from destruc- tor Wood by the very different elytral decliv- ity tliat superficially resembles some male Hylocurus. Male.— Length 2.3 mm (paratypes 1.3-1.4 mm), 2.2 times as long as wide; color very dark brown. Frons and pronotum about as in destructor, except pronotum less scabrous. Elytra proportions and outline as in de- structor; disc similar to destructor except in- terstrial crenulations lower, not as sharp, wider, mostly uniseriate, those on interstriae 1 to 9 near base of declivity largely sub- nodulate but ending at margin of declivity except 9 continuing as a strongly elevated, coarsely serrate, submarginal costa to apex; declivity about as steep but less strongly arched, with interstriae 1 to 8 unarmed, smooth, brightly shining, each with a unise- riate row of minute punctures; vestiture finer and longer than in destructor. Female.— Similar to male except irregu- larly convex. Type material.— The male holotype, fe- male allotype, and three paratypes were taken along the Patzucuaro-Ario de Rosales highway, Michoacan, Mexico, 31-X-1980, 2240 m, S-137, Prunus serotina, T. H. Atkin- son and A. Aquihua. Pseudothysanoes fimbriatus, n. sp. This species is distinguished from peni- culus Wood by the larger size, by the flat female frons, with shorter hair on the ver- tex, and by the different antenna. Female.— Length 1.7 mm (paratypes 1.6-1.9 mm), 2.8 times as long as wide; color dark brown. Frons flat below, very slightly convex above; smooth, shining, and impunctate on median area below, grading into reticulation upward and laterally then finally pimctate- subgranulate; vestiture consisting of a fringe of long hair on upper margin, tips of longest extending slightly beyond middle of frons. Antenna similar to peniculus except segments of scape wider, club slightly narrower; scape, funicle, and club ornamented by rather abun- dant, long setae. Pronotum and elytra as in peniculus except strial pimctures on disc more distinct, inter- strial punctures more nearly replaced by granules. Male.— Similar to female except frons with a large, shallow, central fovea, re- ticulate in lower area, specialized setae on vertex absent, sparse setae uniformly dis- tributed; antenna normal, setae sparse, rather short; pronotum and elytra as in male penic- ulus except dechvital interstriae 3 with a row of rounded granules; elytral vestiture al- most all abraded, apparently similar to male penicidus. Type material.— The female holotype, male allotype, and 18 female paratypes were taken at Zacapoaxtla, Pueblo, Mexico, 6-V- 1981, 2150 m, S-231, Phoradendron, T. H. At- kinson and A. Aquiliua. Pseudothysanoes pini, n. sp. This species is distinguished from the close- ly allied coniferae (Wood) by the less strongly expanded female scape and by differences in the elytral declivity described below. Female.— Length 1.6 mm (paratype 1.5 mm), 3.0 times as long as wide; color black, with white vestiture. Head about as in coniferae except frontal setae less conspicuous; scape less strongly flattened, more nearly like male coniferae with larger tuft of hair. Pronotum and elytral disc about as in com- iferae except interstrial scales closer, shorter, and wider, each scale about as wide as long. Elytral declivity steeper and more broadly 230 Great Basin Naturalist Vol. 42, No. 2 convex than in coniferae; interstriae 2 feebly impressed, 1 and 3 weakly convex on upper two-thirds, weak impression near apex ex- tending from striae 1 to 4; interstriae 5 and 7 feebly elevated, joining, their continuing con- vexity forming ventrolateral margin and fus- ing with 1 at apex; scales on interstriae 1, 3, 5, and 7 forming double rows. Type material.— The female holotype and one slightly crushed female paratype were taken at Km 43 on the Texcoco-Cal- pualpan Highway, Mexico, Mexico, 17-III- 1981, 2780 m, S-198, Pinus hartivegii x P. montezumae, T. H. Atkinson. The holotype and paratype are in my collection. Scolytodes pilifer, n. sp. This species is distinguished from in- gavorus Wood by the much longer setae on the female frons, by the absence of re- ticulation on the pronotum, and by other characters described below. Female.— Length 1.4 mm (male 1.3-1.4 mm), 2.4 times as long as wide; mature color very dark brown. Frons apparently as in ingavonis except setae on upper margin much longer, attain- ing epistoma (greater part of surface ob- scured by these setae). Pronotum as in ingavonis except surfaces between asperities and between punctures smooth, brightly shining, punctures apparent- ly slightly larger, deeper. Elytra as in ingavorus except punctures deeper, strial and interstrial punctures sub- equal in size, those on declivity similar to disc, confused only on lower half, strial setae half as long, interstrial setae stouter. Male.— Similar to female except frons strongly convex, smooth, shining in central area, some reticulation elsewhere, a distinct transverse impression at level of antennal in- sertion, punctures rather fine, not close; ves- titure of fine, short, sparse hair, interstrial bristles on declivity slightly stouter. Type material.— The female holo- type,male allotype, and three paratypes were taken at Uxpanapa, Veracruz, Mexico, 26-V- 1981, 120 m, S-287, A. Aquihua. The holo- type, allotype, and paratypes are in my collection. Scolytus binodus, n. sp. Although this species bears a superficial rsemblance to costeUatus Chapuis, it is not closely related. The unique feature is the presence of a widely separated pair of no- dules on the female sternum 3. The male is unknown. Female.— Length 3.3 mm (paratypes 2.7-4.0 mm), 2.0 times as long as wide; color very dark brown to almost black. Frons broadly convex; surface rather coarsely, convergently aciculate; a few fine punctures in grooves; vestiture of uniformly distributed fine long hair, upper margin on vertex bearing a row of coarser, longer hair, hair on lateral margins from epistoma to up- per level of eyes longer, coarser, more abun- dant. Ventral half of suture 1 of antennal club with a septum, remainder of finely pu- bescent club unmarked by sutures. Pronotum as long as wide; surface smooth, shining; median two-thirds very finely punc- tured, lateral areas more closely, somewhat coarsely punctured; glabrous. Elytra 1.17 times as long as wide, 0.9 times as long as pronotum; surface smooth, shining; posterior third with striae and middle third of interstriae equally, rather deeply impressed (resembling costeUatus), widths and depths of grooves equal on striae and interstriae, these grooves equal in width to convexities separa- ting them from one another, strial and inter- strial punctures in this area about equal; strial grooves continuing to base, interstrial grooves decrease in depth until almost obso- lete at base, punctures decrease in size to half that of striae. A few short, stout interstrial setae on posterior half. Anterior margin of sternum 2 costate, ster- num 2 ascending at rate of about 80 degrees, smooth, shining, a few coarse punctures on its middle third; sternum 3 slightly longer than normal, impunctate, armed by a pair of mod- erately coarse, rounded tubercles, these di- viding segment into approximately equal thirds; sterna 4 and 5 rather finely, closely punctured; a few hairlike setae on sternum 2. Type material.— The female holotype and 10 female paratypes were taken at Uxpa- napa, Oaxaca, 24-V-1981, 120 m, S-182, Combretom sp., A. Equihua. The holotypes and paratypes are in my collection. June 1982 Wood: New American Scolytidae 231 Literature Cited 1982. The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a Wood, S. L. 1974. New species of American bark bee- taxonomic monograph. Great Basin Nat. Mem. ties (Scolytidae: Coleoptera). BYU Sci. Bull., Biol. 6:1-1359. Ser. 19(1); 1-65. TEMPERATURE-RELATED BEHAVIOR OF SOME MIGRANT BIRDS IN THE DESERT George T. Austin' and J. Scott Miller' Abstract.- Behavior of migrant birds in relation to temperature was studied and compared to that of resident species in the northern Mojave Desert. Migrants reduced foraging intensity above 30 C, but resident species showed no striking decrease in intensity of foraging at temperatures up to 35 C. Migrant species shifted activities to shaded microhabitats at temperatures between 20 and 30 C; the resident Verdin showed a similar shift at 35 C. Most mi- grants decreased the amount of time spent foraging at temperatures above 30; Verdins showed a similar but stronger response to temperatures about 30 C. Significant reductions in the use of hovering and hawking maneuvers were found among migrants at temperatures above 30 C. Migrants showed similar types of behavioral adjustments to tem- peratures as did resident desert species, but they responded earlier in the daily temperature cycle. Desert birds ap- pear to correlate their daily activity strongly with temperature, but nondesert species may respond either to temper- ature or time of day. Several studies have shown that resident desert birds react behaviorally to high am- bient temperatures by shifting activity to cooler, shaded microhabitats and reducing the intensity and amount of activity (Smith 1967, Calder 1968, Ricklefs and Hainsworth 1968, Ohmart 1969, Austin 1976, 1978). Sim- ilar reductions in activity at high temper- atures vi^ere found for certain nondesert spe- cies (Ricklefs 1971, Schartz and Zimmerman 1971, Verbeek 1972). It is unknown whether the midday depression in activity from a wide variety of temperature regimens is cor- related with temperatiu-e or with time of day. Among certain desert species, this de- pression in activity is clearly a function of temperature (Ricklefs and Hainsworth 1968, Austin 1976). Migrant birds in the desert, especially in fall, are exposed to temperatures that exceed considerably those encountered either on the breeding or wintering grounds. The potential lethality of desert heat and aridity was docu- mented by Miller and Stebbins (1964). To mi- grate successfully through the desert may re- quire physiological and behavioral adjustments by the species involved. Behav- ioral differences between migrant wood war- blers (Pamlidae) seen at cooler, higher eleva- tions and those seen on the desert floor were noted previously by Austin (1970). This study was designed to detect and quantify behav- ioral changes over a range of temperatures by small passerine migrants during fall in the northern Mojave Desert. Methods Areas frequented by migrants were visited periodically throughout the peak migration period from 20 August to 25 September 1975. These included sites near Las Vegas, Tule Springs Park, Com Creek Field Station of the Desert National Wildlife Range, and Moapa Valley, all in Clark Co., Nevada, and at Beaver Dam Wash, Mohave Co., Arizona. Most data on migrants were obtained at Tule Springs and Com Creek, where numerous de- ciduous trees have been planted. Data on resident species and on a few migrants were obtained in natural desert riparian vegetation dominated by mesquite {Prosopis juliflora) near Las Vegas and at Corn Creek. When a bird was encountered, the follow- ing data were recorded: species of bird; amount of time spent in each of several ac- tivities (sitting, preening, flying, foraging), timed with a stopwatch; amount of time spent in either shade or sun; number of perch changes per stopwatch-timed interval; num- ber of each foraging maneuver (glean, hover, hawk as described by Root 1967); time of day 'Nevada State Museum, Capitol Complex, Carson City, Nevada 89710. 232 June 1982 Austin, Miller: Migrant Desert Birds 233 and air shade temperature (T/s noted on the half h). Several problems were encountered by the observer while following the individ- ual bird under observation; the most notable of these was keeping track of the individual, which was often among several conspecifics. This situation, and the fact that migrants rarely remained in sight for long periods, re- sulted in most observations being of 3-5 min- utes or less duration. Mean values and per- centages were calculated using all periods of observation regardless of duration. Only ob- servations greater than 1 minute in length were subjected to further statistical analysis. Foraging maneuver data were treated with chi-square goodness of fit on the numerical data using Yates's correction where appli- cable. Feeding activity as perch changes per min and microhabitat usage expressed as pro- portion of time (transformed to arcsine; Zar 1974) spent in shade were analyzed using polynomial regression analysis. Diversity of foraging maneuvers and partitioning of time for T/s above and below 30 C were calcu- lated using the information-theoretical mea- sure to the base e (H'). Differences between H' at the two temperature ranges were tested using the methods outlined in Zar (1974). Sta- tistical significance was set at P < 0.05 throughout. Principal species studied were (minimum number of individuals observed are in paren- theses): Warbling Vireo (Vireo gilviis) (20) and the warblers: Orange-crowned (Verrni- vora celata) (32), Nashville (V. nificapilla) (25), Yellow {Dendroica petechia) (68), Mac- Gillivray's {Oporonis tolmiei) (30), Wilson's {Wilsonia pusilla) (37), and American Red- start {Setophaga ruticilla) (7). Additional small samples (less than 50 minutes and 10 in- dividuals) were obtained for other migrant and some resident species that will be men- tioned in the text. Data were also obtained for the resident Verdin {Auriparus flaviceps); these were combined with data gathered for other studies (Austin 1976, 1978). Total sample for Verdins was approximately 30 in- dividuals. Total amount of timed data for each temperature range is given in Table 1. Data collection was facilitated by the use of tape recorders. Observations were made on clear days under low wind conditions (< 8 km/h). Results Foraging rate.— Rate of foraging by mi- grant species was comparatively rapid at T^'s below 30 C and sharply less rapid above 30 C (Table 2). On the average, foraging by mi- grant warblers was reduced by 10 perch changes per minute (37 percent) above 30 C. Greatest reductions were by Orange-crowned and Nashville warblers (43 and 47 percent). Analysis of these data by 2 C intervals in- dicated that 30 C was a well-defined point where foraging rate changed abruptly. Rate of foraging by Yellow Warblers, for example, average 22.1 and 15.2 perch changes per minute at 28-30 C and 30-32 C, respectively. Table 1. Amount of observation time (seconds) obtained for foraging behavior. Ta (degrees C) 15-20 20-25 25-30 30-35 35-40 Verdin {Vf 10576 49475 45813 41179 59800^' Bewick's Wren (BW) - 403 594 1235 - Black-t. Gnatcatcher (BTG) — 677 460 — 201 Warbling Vireo (WV) 1039 1732 1324 1283 — Orange-cr. Warbler (OCW) 1883 1621 3583 2371 — Nashville Warbler (NW) 1238 1409 1050 3357 1443 Virginia's Warbler (VW) 372 772 85 426 180 Lucy's Warbler (LW) 50 789 67 1023 — Yellow Warbler (YW) 3179 5783 8307 16475 1327 Black-th. Gray Warbler (BTGW) 132 538 48 793 — Townsend's Warbler (TW) 1966 — 344 698 — MacGillivray's Warbler (MW) 2323 2928 1006 3181 681 Wilson's Warbler (WW) 2071 1341 4688 5259 370 American Redstart (AR) 657 533 801 593 — ^Species code in parentheses. Wa = 35-50 degrees C. 234 Great Basin Naturalist Vol. 42, No. 2 Apparently this response was more a func- tion of Ta than time of day as indicated in Table 3. Rate of foraging during midday was greater on cool days than on hot days. Early morning foraging was at a somewhat reduced rate compared to later in the day at similar Tg's, accounting for the lower intensities at 15-20 C. Because of this, data for temper- atures below 20 C were not used in regres- sion analysis (below). The slowly foraging Warbling Vireo showed only a slight decrease in foraging above 30 C. Its foraging rate, however, was lower than other species at all T^'s. Among migrant species with small samples, Virginia's Warbler (Vertnivora virginiae) exhibited a gradual decrease in foraging with increasing T^. American Redstart decreased foraging rate dramatically (by 44 percent) at 30 C and Townsend's Warbler {Dendroica townsendi) decreased foraging at about 25 C. The mi- grant Black-throated Gray Warbler (D. ni- grescens) and the resident Bewick's Wren {Thnjornanes bewickii). Black-tailed Gnatcat- cher {Polioptila melanura), and Lucy's War- bler (V. luciae) apparently do not reduce fo- raging through at least 35 C. The Verdin exhibited a gradual decrease in rate of forag- ing with increasing T^ (Table 2; Austin 1976: Fig. 6); the most dramatic decrease (45 per- cent) was at 35 C. Although the individual observations show considerable variability at all T^'s, foraging rates of the various species exhibit significant negative linear or quadratic relationships with Tj, (Table 4). Although the variability in- volved is real on a short-term basis, we be- lieve that the averages for each temperature range (Table 2) reflect a true and biologically important adjustment by these birds as T^ in- creases. Samples obtained over longer peri- ods of time on an individual tend to show less variability, as indicated previously for the Verdin (Austin 1976). MicROHABiTAT USAGE.— Migrant species varied in the relative amount of time spent in shaded or exposed microhabitats (Fig. 1). MacGillivray's and Wilson's warblers foraged largely in the shade at all T^'s but especially at higher T^'s. In other transients, the propor- tion of time spent foraging in the shade was not as great at lower T^'s and increased rap- idly with T^. Warbling Vireos and Yellow Warblers made the most abrupt shift to shaded microhabitats at 30 C; Orange- crowned and Nashville warblers did so at 20 C. On the average, more than 95 percent of all foraging was in shade at T^'s above 30 C (Fig. 1). The resident Verdin, in contrast, did not shift its foraging to predominately shaded mi- crohabitats until Ta exceeded 35 C (Fig. 1; Austin 1976: Fig. 5). Above 35 C, about 15 percent of its foraging was still in vegetation exposed to the sun. Data for the Black-tailed Gnatcatcher indicated a similar pattern. Bewick's Wren and Lucy's Warbler appeared to forage predominately in the shade at all Ta's. As with foraging rate, individual varia- bility in microhabitat use (as proportion of time spent in shade) was great. Regression analysis of time spent in shade in relation to Ta indicated significant positive correlations for the six migrant species (Table 4). The var- iability and consequently low correlations for MacGillivray's and Wilson's warblers were likely due to the large number of relatively Table 2. Mean number of perch changes per minute by birds in relation to ambient temperature.' Ambient temperature (degrees C) Species 15-20 20-25 25-30 30-35 35-40 22.0 18.2 19.4 16.6 9.1 15.4 16.4 14.8 11.2 — 21.6 26.2 23.0 13.1 — 18.6 21.5 26.1 13.9 10.8 22.4 22.4 23.2 13.5 13.4 20.6 26.7 21.2 15.1 — 32.7 37.0 34.6 26.6 19.8 31.6 41.1 39.5 23.3 — Verdin Warbhng Vireo Orange-crowned Warbler Nashville Warbler Yellow Warbler MacGillivray's Warbler Wilson's Warbler American Redstart Observations summed for each temperature range regardless of duration of each observation; total observation time for each temperature range as in- dicated in Table 1. June 1982 Austin, Miller: Migrant Desert Birds 235 short-duration (1-2 min) observations for these species. The statistically significant regressions, however, indicated that all six species increased the use of shade as T^ increased. Time budget.— The partitioning of time by migrants varied with T^. At lower T^'s, nearly all time was spent foraging and less than 5 percent spent sitting (Fig. 2). Above 30 C, the amount of time sitting increased to about 20 percent in most species. MacGil- hvray's Warbler did not increase time spent in inactivity above 20 percent until T^ reached 35 C. Wilson's Warbler did not sit more than 6 percent of the time at any T^. The Verdin increased sitting time gradu- ally through 35 C. Above 35 C, the amount of time sitting increased sharply to 66 per- cent (Fig. 2; Austin 1978). The other three resident species did not increase time spent sitting above 30 C; too few data were ob- tained for T/s above 35 C to draw conclusions. Foraging maneuvers.— At T^'s greater than 30 C, all migrant species decreased the use of hawking and hovering foraging ma- neuvers (Fig. 3). This change in foraging strategy above and below 30 C was statis- tically significant in all species except Black- throated Gray and MacGillivray's warblers I I EXPOSED MICROHABITAT rZ<^ SHADED MICROHABITAT 12345 12345 T W M W ■ 12 3 4 5 12 5 4 5 12 3 4 5 12 3 4 5 12 3 4 5 AMBIENT TEMPERATURE Fig. 1. Percent of time spent in shaded and exposed microhabitats as a function of ambient temperature by mi- grant and resident birds in southern Nevada. Temperature code is as follows: (1) 15-20 C, (2) 20-25 C, (3) 25-30 C, (4) 30-35 C, (5) 35-40 C. Species indicated by letters above bars are as coded in Table 1. 236 Great Basin Naturalist Vol. 42, No. 2 and was especially striking in Wilson's War- bler and American Redstart. Overall, both hawking and hovering were reduced propor- tionately, but Wilson's Warbler reduced hov- ering to a greater extent than hawking. When all three types of maneuvers were con- sidered independently. Warbling Vireo, Yel- low and Wilson's warblers, and American Redstart exhibited significant changes in overall foraging behavior. Larger samples for other species may show similar differences. Samples for any one of the resident species were too small for analysis. The pooled data for the four species, however, showed no change in foraging strategy over the range of Tj's sampled. Discussion Migrant species of birds showed behavioral changes at high temperatures that were sim- ilar in kind to those of resident species. Both reduced the amoimt and rate of foraging and shifted their activities to cooler microhabitats as ambient temperatures increased. Such changes in behavior reduce metabolic and environmental heat gain. In migrants the shift to cooler microhabitats tended to pre- cede reduction in foraging rate, as was pre- viously found for residents (Ricklefs and Hainsworth 1968, Austin 1976). This allows reduced heat stress but continued high rates of foraging. Changes in behavior were appar- ently more closely related to temperature than to time of day. Although migrants were similar to resident desert species in the types of behavioral changes with increasing temperature, there were obvious differences in the temperature at which these occurred and the magnitude of change. Among resident species, foraging was reduced only at T^'s exceeding 35 C. Mi- grants generally decreased foraging rates at about 30 C. Similarly, there was an abrupt change in the partitioning of time by mi- grants at 30 C and by residents at 35 C. A shift to shaded microhabitats occurred in mi- grants at Ta's ranging between 20 and 30 C. Residents did not greatly increase the use of shade until T^'s exceeded 35 C. At T^'s exceeding 35 C, the resident Verdin drastically reduced the amount of time spent foraging to less than 35 percent. No migrant species decreased foraging to less than 70 percent. This suggests that migrants may have to continue foraging at high T^'s to re- plenish energy stores and to maintain water balance, or that their adjustments to T^ are less refined than those of residents. The mi- grants' unfamiliarity with local conditions may also, in part, account for the increased foraging time. In any case, it seems likely that migrants need access to free water or an abundant food source during passage through the desert, which accounts for their relative rareness in the low desert away from riparian habitats. Table 5 shows that diversity of foraging maneuvers decreased in all species at T^'s above 30 C; these differences were signifi- cant for all species except MacGillivray's Warbler. This decrease in diversity reflects the reduction of maneuvers involving flight (Fig. 3). Diversity in the partitioning of time increased significantly at T^'s above 30 C (Table 5). This reflected the increased amount of time spent resting at higher T^'s, whereas nearly all time was occupied by fo- raging at the lower T^'s (Fig. 2). Extreme weather conditions were shown by Grubb (1975) to affect foraging diversity of birds in eastern deciduous forest. Diurnal rhythms in foraging behavior have long been recognized in birds (e.g., Palmgren 1949). It is generally recognized that there is a peak of activity in the morning that Table 3. Foraging rate (number of perch changes per minute) as a fimction of T^ and time of day. Time 0600-1000 1000-1400 1400-1800 Species T^ < 30 C < 30 C > 30 C >30C Orange-crowned Warbler Nashville \\{^rbler Yellow Warbler MacGillivray s Warbler Wilson's Warbler 22.9 24.2 14.3 12.5 19.4 23.1 14.0 12.8 21.5 27.3 16.9 13.8 22.3 26.2 15.1 — 34.2 35.1 26.0 17.4 June 1982 Austin, Miller: Migrant Desert Birds 237 decreases toward midday and often increases again to another peak at the end of the day. In some species, this appears as a circadian rhythm, occurring at the same time of day under constant hght conditions, as in the Arc- tic (Armstrong 1954) and under constant tem- perature conditions in the laboratory (Eyster 1954, Coutlee 1968, Smith et al. 1969). High temperatures modified this behavior in at least the White-crowned Sparrow (Zono- trichia leucophrys); activity at 33 C was 50 percent of that at 23 C. Daily activities of the House Sparrow {Passer domesticus) and Dark-eyed Junco (Junco hyemalis), however, were affected little at temperatures as high as 32 C and 35 C, respectively (Eyster 1954). Field studies have demonstrated that a number of nondesert bird species reduce ac- tivity at high temperature. In the post- breeding Yellow-billed Magpie {Pica nut- talli), the amount of time spent foraging was negatively correlated with temperatures above approximately 25 C (Verbeek 1972). The male Dickcissel {Spiza americana) de- creased various behaviors associated with re- production and increased the amount of time spent resting as temperatures increased above 35 C (Schartz and Zimmerman 1971). Al- though total time spent foraging seemed unaffected, foraging intensity was reduced. The midday decrease in foraging time by the tropical Mangrove Swallow {Iridoprocne alhi- linea) was also suggested to be due to high radiational heat loading coupled with high humidity (Ricklefs 1971). Tropical fly- catchers and Temperate zone swallows showed little or no reduction in midday ac- tivities (Ricklefs 1971). Several desert species showed a close cor- relation between a reduced rate of activity or increased use of shaded microhabitats and in- creasing temperature (Dawson 1954, Smith 1967, Calder 1968, Ricklefs and Hainsworth 1968, Ohmart 1969, 1973, Austin 1976, 1978). Most species are highly active at mid- day in the cooler months of the year or on cool, cloudy days during midsummer. Behav- ioral changes observed in these studies may be actual adjustments in direct response to Ta, an indirect reflection of changes in distri- bution and activity of prey items or a com- bination of them. The available data suggest the latter. The Roadnmner {Geococcys cali- fornianus) maintained approximately con- stant rates of activity throughout the day in the laboratory at moderate temperatures (Ka- vanau and Ramos 1970). In the field, well- defined periods of inactivity in shaded areas were apparent during hot middays (Calder 1968, Ohmart 1973). Time of foraging was correlated with peak activity of their princi- pal prey (Ohmart 1973). Captive Cactus Wrens {Campylorhynchus brunneicapillus) decreased activity and increased amount of time spent in shade with increasing T^ with food equally available under all conditions (Ricklefs and Hainsworth 1968). These changes in behavior were similar to those Table 4. Regression analyses of the relationships between ambient temperature (X = Tab) ^"d foraging intensity (Y = perch changes per minf and proportion of time spent in shaded microhabitat (Y = arcsine of the proportion) by some migrant birds in the Mojave Desert. Dependent variable Number of Correlation Regression Species (Y) observations^ coefficient formula Warbling Vireo perch changes/min 25 0.477 Y = 29.20 - 0.53X % of time in shade'' 21 0.558 Y = -27.70 + 3.19X Orange-crowned Warbler perch changes/min 33 0.695 Y = 57.98 - 1.27X % of time in shade'' 35 0.652 Y = -22.81 + 2.98X Nashville Warbler perch changes/min 42 0.646 Y = -155.97 -t- 13.01X - 0.24)^ % of time in shade'' 25 0.696 Y = -6.45 + 2.71X Yellow Warbler perch changes/min 100 0.500 Y = 43.41 - 0.82X % of time in shade'' 81 0.661 Y = 188.01 - 13.20X -1- 0.30X2 MacGillivray's Warbler perch changes/min 36 0.514 Y = 45.87 - 0.89X % of time in shade'' 33 0.351 Y = 60.29 + 0.75X Wilson's Warbler perch changes/min 71 0.467 Y = 66.47 - 1.20X % of time in shade'' 41 0.442 Y = 25.68 + 1.65X ^nly data for ambient temperatures greater than 20 C were used, because foraging intensity was reduced in early morning al lower temperatures. Relationship expressed as arcsine Y = arcsine a + bX. '^Only observations > 1 min in length were used in regression analysis (see text). 238 Great Basin Naturalist Vol. 42, No. 2 observed in the field. Data on insects indicate that they reduce their activity and move into shaded microhabitats in response to increas- ing temperature (e.g., Clench 1966, Austin 1977). Aside from differences in prey distribution and activity and temperature, other factors may account for some of the variability ob- served in behavioral modifications. The amount of time spent resting by the Dickcis- sel increased more rapidly at high T^'s when relative humidities were greater (Schartz and Zimmerman 1971). In the Las Vegas area during August and September, relative hu- midities average below 20 percent during daylight hours (Brown 1960) and are unlikely to be a major factor affecting behavior. Wind also affects bird behavior (Grubb 1975). In this study, however, observations were not made when the wind exceeded 8 km/h (most were taken on completely windless days), so wind is considered a constant. Two distinct thermoregulatory problems arise when contending with desert heat; heat loading is often severe and, although low hu- midities promote effective evaporative cool- ing, water is largely unavailable. Species oc- curring in the desert are thus faced with opposing problems of water conservation and maintenance of heat balance. Such behav- ioral adjustments as shifting to shaded micro- habitats and eventually reducing the intensity I 2 3 4 S WV m^ rm D 12 3 4 5 OC W I 2 3 4 S N W I 2 3 4 S V W bcbK^^ Hm^ •2345 12345 12345 12345 2 3 4 5 12 3 4 5 ra& 72B^ 12 3 4 6 12 3 4 5 2 3 4 5 AMBtENT TEMPERATURE Fig. 2. Time budget of migrant and resident birds as a function of ambient temperature in southern Nevada. Tem- perature and species code as in Fig 1. June 1982 Austin, Miller: Migrant Desert Birds 239 and amount of activity to a minimum may be a necessary and important means of con- tending with high temperatures. In non-arid localities behavioral means of reducing heat load may also be of some importance but for a different reason. Although temperatures tend to be lower than in arid areas, relative humidities are usually higher. This increased humidity reduces the effectiveness of evapo- rative cooling mechanisms due to a decrease in the vapor pressure gradient. Further stud- ies of bird behavior under varying humidity conditions are highly desirable. Acknowledgments We thank W. G. Bradley, K. S. Moor, S. Naegle, R. D. Ohmart, R. E. Ricklefs, D. Thomas, and J. L. Zimmerman for sugges- tions on statistical procedures or comments and suggestions for improvements of the manuscript. Literature Cited Armstrong, E. A. 1954. The behavior of birds in contin- uous dayhght. Ibis 96:1-30. Austin, G. T. 1970. Migration of warblers in southern Nevada. Southwest. Natur. 15:2.31-237. 1976. Behavioral adaptions of the Verdin to the desert. Auk 93:245-262. 1977. Notes on the behavior of Asterocampa leilia (Nymphalidae) in southern Arizona. J. Lepid. Soc. 31:111-118. 1978. Daily time budget of the post-nesting Ver- din. Auk 95:247-251. Brown, M. 1960. Climates of the states, Nevada. Cli- matography of the United States. U.S. Dept. of Commerce, Weather Bureau, No. 60-26. Calder, W. a. 1968. The diurnal activity of the Road- runner, Geococcys californianus. Condor 70:84-85. Clench, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology 47:1021-1034. CouTLEE, E. L. 1968. Maintenance behavior of Lesser and Lawrence's goldfinches. Condor 70:378-384. Dawson, W. R. 1954. Temperature regulation and wa- ter requirements of Brown and Albert towhees, Pipilo fiiscus and Pipilo aberti. Univ. of California Publ. Zool. 59:81-124. w 242 71 OC W 535 117 N 251 W 213 Y 1156 W 623 80 - 60- 40 — 20 - n — ll ^ ^ D GLEAN M AWK HOVER 00—1 80- BT 24 OW 42 T 125 W 69 M 255 W 175 W 3S0 w 608 A M 2 R 32 60- 40- 20 — o — 1 ^ ^^ ^^ m ^g P ^^ Fig. 3. Changes in the use of various foraging maneuvers as a function of ambient temperature by migrant birds in southern Nevada. Left-hand bar for each species represents temperatures less than 30 C; right-hand bar represents temperatures greater than 30 C. Species code as in Fig. 1. Number of foraging maneuvers observed indicated above each bar. 240 Great Basin Naturalist Vol. 42, No. 2 Table 5. Indices of foraging diversity (H') by migrant birds in relation to temperature. Maximum diversity for 3 alternatives = 1.099 (to the base e). Time budget at T^ Foraging maneuvers at Ta Species <30C > 30 C < 30C >30C Warbling Vireo 0.087 0.263 0.621 0.290 Orange-crowned Warbler 0.219 0.539 0.359 0.136 Nashville Warbler 0.340 0.584 0.376 0.185 Yellow Warbler 0.200 0.418 0.329 0.157 MacGillivray's Warbler 0.150 0.419 0.177 0.098* Wilson's Warbler 0.080 0.256 0.961 0.567 American Redstart 0.112 0.661 0.949 0.371 ^Difference not significant. Eyster, M. B. 1954. Quantitative measurement of the influence of photoperiod, temperature, and sea- son on the activity of captive songbirds, Ecol. Monogr. 24:1-28. Grubb, T. C, Jr. 1975. Weather-dependent foraging be- havior of some birds wintering in a deciduous woodland. Condor 77:175-182. Kavanau, J. L., AND J. Ramos. 1970. Roadrunners: activ- ity of captive individuals. Science 169:140-147. Miller, A. H., and R. C. Stebbins. 1964. The lives of desert animals in Joshua Tree National Mon- ument. Los Angeles, Univ. of California Press. Ohmart, R. D. 1969. Physiological and ethological adap- tations of the Rufous-winged Sparrow {Aimophila carpalis) to a desert environment. Unpublished dissertation. Univ. of Arizona. 1973. Observations on the breeding adaptations of the Roadrunner. Condor 75:140-149. Palmgren, p. 1949. On the diurnal rhythm of activity and rest in birds. Ibis 91:561-576. RiCKLEFS, R. E. 1971. Foraging behavior of Mangrove Swallows at Barro Colorado Island. Auk 88:635-651. RiCKLEFS, R. E., and R. R. Hainsworth. 1968. Temper- ature dependent behavior of the Cactus Wren. Ecology 49:227-233. Root, R. B. 1967. The niche exploitation pattern of the Blue-gray Gnatcatcher. Ecol. Monogr. 37:317-350. ScHARTZ, R. L., and J. L. Zimmerman. 1971. The time and energy budget of the male Dickcissel {Spiza americana). Condor 73:65-76. Smith, E. L. 1967. Behavioral adaptations related to wa- ter retention in the Black-tailed Gnatcatcher (Po- lioptila melanura). Unpublished thesis. Univ. of Arizona. Smith, R. W., I. L. Brown, and L. R. Mewaldt. 1969. Annual activity patterns of caged non-migratory White-crowned Sparrows. Wilson Bull. 81:419-440. Verbeek, N. a. M. 1972. Daily and annual time budget of the Yellow-billed Magpie. Auk 89:567-582. Zar, J. H. 1974. Biostatistical analysis. Englewood Cliffs, New Jersey, Prentice-Hall, Inc. DESCRIPTION OF A NEW PHALACROPSYLLA AND NOTES ON P. ALLOS (SIPHON APTERA: HYSTRICHOPSYLLIDAE) R. B. Eads' and E. G. Campos' Abstract.- PhalacropsijUa morlani, from New Mexico is described as new to science and figured. It is separable from the other species in the genus by the possession of 18 spines in the pronotal comb and by the distinctive shape and setation of the male distal arm of sternum IX. Other species in the genus have 14 to 16 spines in the pronotal comb. Host and distributional records for P. alios are given. The description of Phalacropsylla morlani brings to six the number of known species in this montane genus. Other species are P. par- adisea Rothschild 1915, P. alios Wagner 1936, P. nivalis Barrera & Traub 1967, P. Jiamata Tipton & Mendez 1968, and P. orego- nensis Lewis & Maser 1978. Normal hosts of the Phalacropsylla are believed to be wood rats, Neotoma spp., and closely associated ro- dents and lagomorphs. A key is provided to aid in separating the species. Phalacropsylla morlani, sp.n. (Figs. 1-2) Type material.— Holotype male ex Och- otona princeps (Richardson), Santa Fe, New Mexico, 10 Nov. 1958, H. B. Morlan, eleva- tion ca 3048 m (10,000 ft). Morlan (pers. comm.) reports that a second male of this species with same collection data has been lost. Diagnosis.— Phalacropsylla morlani most closely resembles P. alios and P. nivalis in that there are no long, curved spiniforms pre- apically on the outer surface of the male st IX, as is the case with P. oregonensis, P. ham- ata, and P. paradisea. In P. nivalis there is a deep sinus in the caudal margin of the male fixed process. In P. alios there is a shallow sinus in the caudal margin of the male fixed process, but the margin is merely sinuate in P. morlani. The pronotal comb of P. morlani has 18 teeth; there are 16 in P. alios. Description of male.— Head (Fig. 1): Preantennal region with 2 slightly concave rows of bristles. Frontal row of 4 small, thin bristles with 3 fine intercalaries; ocular row of 4 much longer bristles; ca 5 thin bristles caudad of the ocular row. Maxilla narrow, acuminate distally extending to base of 4th segment of maxillary palpus; maxillary pal- pus extending almost to apex of coxa I. The 5-segmented labial palpus reaching beyond midportion of trochanter I. Postantennal re- gion with bristles arranged 1-3-6 on one side, 2-3-6 on other, the caudal row with fine in- tercalaries; 16 or 17 fine hairs in an irregular row along the antennal fossa. Thorax: Pronotum with a row of 7 large bristles separated by about same number of smaller ones per side; pronotal comb of 20 spines, all of approximately same length ex- cept for small ventralmost pair; second ven- tralmost pair appreciably wider than others. Mesonotum with a row of 6 large bristles and an equal number of fine intercalaries preced- ed by a row of 5 or 6 smaller ones and 15 to 20 short bristles scattered on cephalad mar- gin; mesonotal flange with 3 pseudosetae per side. Mesepisternum with 2 subequal lateral bristles. Mesepimeron with 2 irregular rows of 2 bristles, caudal row of 6 or 7 long bristles with fine intercalaries, cephalad row of 5 or 6 smaller bristles. Metepisternum with a long bristle on subdorsal margin and a short one on the dorsal margin. Metepimeron with ca 5 lateral bristles arranged 2:2:1. 'Vector-Bome Viral Diseases Division, Center for Infectious Diseases, Centers for Disease Control, Public Health Service, U.S. Department of Health and Human Services, P.O. Box 2087, Fort Collins, Colorado 80522-2087. 241 242 Great Basin Naturalist Vol. 42, No. 2 Legs: Procoxa well provided with subequal bristles; dorsal Vz of anterior margin with row of short bristles; ventral Vi with 2 widely spaced long bristles, the ventral bristle reach- ing ca V2 length of profemur: subventral row of 6 long bristles extend well beyond trochan- ter. Profemur with row of short bristles on anterior margin, 2 submarginal bristles at dis- tal third, and 2 irregular rows of ca 15 long- er, thin, lateral bristles extending length of femur. Metacoxa outer surface without bristles on upper third, lower % with scat- tered bristles of unequal sizes, and an oblique row of 3 large bristles near ventral margin extending well below apex of trochanter; 3 large bristles near apex; inner surface of metacoxa with thin setae along anterior mar- gin and submarginal, widening apically to in- clude an oblique row of about 8 small spin- iform bristles. Mesocoxa bristles on inner side limited to anterior margin with submarginal bristles on apical half becoming progressively larger toward apex; 3 large median bristles on ventral margin. Metatibia with 3 bristles, 2 long and 1 short at apex of dorsocaudal margin, the longest of which extends well beyond first tarsal segment; above these bristles are 6 notches bearing subequal, stout bristles, from apex to base 2:3:2:2:2:1. Abdomen: Terga I-IV with apical spine- lets: (1-0), (1-1), (1-1), and (1-0). Terga typi- cally with a row of small bristles (1-5), fol- lowed by a row of larger bristles (4-7), alternating with smaller ones. Unmodified sterna with a vertical row of 1 to 3 bristles preceded by 1 or 2 smaller ones. Middle an- tepygidial bristle ca 2 X length of ventral bristle and almost 3 X length of dorsal bristle. Modified abdominal segments: St VIII roughly triangular, higher than greatest width, devoid of bristles except for 1 large Fig. 1. Phalacropsylla morlani, male: A, head, prothorax and procoxa; B, clasper June 1982 Eads, Campos: A New Flea 243 and 2 small bristles near midpoint of ventral margin; anterior margin fairly straight, even- ly rounded at juncture with dorsal margin; dorsal and ventral margins join at blunt, evenly rounded, caudal apex. Immovable process large and well provided with bristles, especially on dorsal margin; caudal margin with 5 large bristles toward bluntly rounded jvmcture with dorsal margin; caudal margin sinuate but without pronounced sinus divid- ing process into lobes. Movable process of clasper ca 5 X as high as width at infra- foveal region, candle shaped, apex reaching almost as high as immovable process; anterior margin fairly straight, with a few scattered, marginal to submarginal setae; distal half of posterior margin fairly straight, basal half convex; thin bristles along most of the poste- rior margin, thickest on basal %. Manubrum long and slender, about 20 X as long as broad at midpoint; anterior margin straight, posterior margin convex at midpoint. St IX V-shaped, proximal arm much shorter than distal arm, and fish tailed at apex as with other species in the genus; distal arm ca 4 X as long as greatest width, a crescentic row of 8 subequal, spiniform setae toward apex, followed by a marginal row of thin bristles extending ca % distance to base. Lightly sclerotized dorsal expansion of distal arm not discernible in holotype. Discussion Within the genus, P. alios has been recov- ered most frequently and from the widest geographical range. Described from speci- mens off Neotoma cinerea, Logan, Utah, it has subsequently been reported from Califor- nia, Montana, New Mexico, and Wyoming. We have taken alios from March to August central sclerite medan dorsal lobe Fig. 2. Phalacropsylla spp., males: C, morlani aedeagus; D, morkini IX sternum; E, alios IX sternum. 244 Great Basin Naturalist Vol. 42, No. 2 in Larimer Co., Colorado, during year-round shrub on the rocky slopes is mountain mahog- rodent trapping on the Weaver Ranch in any, Cercocarpus montanus. 1977 as follows: 1 male ex N. mexicana 6 Stark and Kinney (1969) have reported the Aug.; 1 female ex Reithrodontomys megalotis recovery of 49 P. alios from California from 8 May; 1 male ex Peromyscus difficilis 5 25 N. cinerea nests and 17 from 13 N. cinerea March; 1 female ex P. difficilis 4 April; 1 from shallow caves in the Lava Beds National male ex P. difficilis 8 May; 1 male ex P. diffi- Monument, Siskiyou Co. None were taken cilis 9 May; 1 female ex Peromyscus manicu- from 9 surface nests or 6 N. cinerea trapped latus 7 May; 1 male ex P. maniculatus 7 Aug. on the surface. A single specimen was recov- The Weaver Ranch is 20 km N of Ft. Col- ered from a P. maniculatus obtained at lins, Colorado, on U.S. 287. The relatively ground level. More recently. Dr. B. C. Nelson treeless, foothills habitat varies in elevation (pers. comm.) has collected alios in numbers from ca 1600 m on the prairie to 1900 m at from N. cinerea nests in the same caves: 21 the highest point on the ridge, bisecting the males, 15 females, 9 Dec. 1976; and 1 male, ranch from north to south. The dominant 31 Jan. 1980. Key to the species of Phalacropsylla (Female of morlani unknown) 1. Male 2 — Female 7 2. Fixed process of clasper divided caudally by a deep sinus 3 — Fixed process of clasper without pronounced sinus, posterior margin sinuate 4 3. Two long, curved spiniforms present preapically on inner surface of distal arm of St IX hamata — No long, curved spiniforms on inner surface of distal arm of st IX nivalis 4. Long, curved spiniforms present preapically on inner surface of distal arm of St IX 5 — Without long, curved spiniforms preapically on inner surface of distal arm of St IX 6 5. Apex of movable process extending about to apex of fixed process paradisea — Apex of movable process extending less than % height of fixed process .... oregonensis 6. 16 teeth in pronotal comb alios — 20 teeth in pronotal comb morlani 7. Caudal lobe of st VII longer than broad 8 — Caudal lobe of st VII broader than long 9 8. Caudal lobe of st VII ca 1.5 X as long as broad nivalis — Caudal lobe of st VII ca 1.9 X as long as broad alios 9. Caudal lobe of st VII 3.5 X as broad as long hamata — Caudal lobe of st VII less than 2 X as broad as long 10 10. Caudal lobe of st VII rectangular, broadly rounded at apex paradisea — Caudal lobe of st VII more triangular, apex bluntly pointed and deflected ventrally oregonensis Acknowledgments USPHS (Ret.), who has contributed greatly to our knowledge of vector-borne diseases. Gary This species is named for the collector, O. Maupin, VBVDD, and W. S. Archibald, Harvey B. Morlan, sanitarian director, formerly of VBVDD, were involved in the June 1982 Eads, Campos: A New Flea 245 collection of the P. alios in Larimer Co., Col- orado. Dr. R. E. Lewis, Iowa State Univer- sity, was consulted concerning the taxonomic status of P. morlani. Literature Cited Barrera, a., and R. Traub. 1967. Phalacropsylla ni- valis, a new species of flea from Mexico (Si- phonaptera: Hystrichopsyllidae). An. Esc. Nac. Cienc. Biol. Mexico City 14: 35-46. Lewis, R. E., and C. Maser. 1978. Phalacropsylla orego- nensis sp. n., with a key to the species of Phala- cropsylla Rothschild 1915 (Siphonaptera: Hys- trichopsyllidae). J. Parasit. 64(1): 147-150. Rothschild, N. C. 1915. On Neopsylla and some allied genera of Siphonaptera. Ectoparasites I: 30-44. Stark, H. E., and A. R. Kinney. 1969. Abundance of ro- dents and fleas as related to plague in Lava Beds National Monument, California. J. Med. Ent. 6(3): 287-294. Tipton, V. J., and E. Mendez. 1968. New species of fleas (Siphonaptera) from Cero Potosi, Mexico, with notes on ecology and host parasite relation- ships. Pacific Insects 10(1): 177-214. Wagner, J. 1936. Neue Nordamerikanische Floharten. Z. Parasitenkd. 8; 654-658. VEGETATION OF THE MIMA MOUNDS OF KALSOW PRAIRIE, IOWA Jack D. Brotherson' Abstract.- One hundred and twenty-eight niima mounds were studied relative to their vegetational relationships in a tall grass prairie area of central Iowa. Mound origins are thought to be due to several phenomena but are most likely initiated and maintained by the activity of pocket gophers. Seventy-five percent of the plant species common to the mounds are prairie species. When vegetative composition of the mounds was compared to the adjacent prairie vegetation, however, they were only 35 percent similar. The mounds were shown to alter the original structure and composition of the prairie vegetation. The mounds, once formed, created a new microenvironment. Many species were shown to respond to this new habitat. The factors deemed most influential in affecting the vegetational changes were disturbance and microrelief. Study observations indicate that the mounds represent microsuccession sites and cause changes in prairie vegetation to earlier stages in the sere. Provision for state-owned prairies in Iowa was made in 1933 when the State Con- servation Commission prepared a report known as the Iowa Twenty-five Year Con- servation Plan. In a section of that report the following proposal was recommended: Prairie Preserve— Recommended. Along the railroad rights-of-way, and here and there in small patches throughout the state, unbroken virgin prairie sod is still to be found. Some of these will be saved because they lie within protected areas, or simply because the ground cannot be used for farm purposes. But somewhere in Iowa a large enough original tract of prairie vegetation should be secured in order to save, under control of the state, the characteristic landscape, wild flowers, and wild life of the native prairies. Several tracts ranging from forty to three hundred acres have been found by the survey. The Conservation Plan includes a Prairie Preserve which will be one of the remaining original areas, or which may be produced by purchase of semi- waste land and bringing it back to prairie condition in a few years' time. (Hayden 1945) Prairies now owned by the state of Iowa were purchased and set aside as natural areas with the intent that features typical of prairie landscapes, such as wild flowers and wildlife native to the tall-grass prairie region, could be preserved for posterity. It was also in- tended that these areas would be useful as game and wildlife sanctuaries; as examples of the native prairie soil types, where com- parisons could be made with cultivated soils of the same soil association; and as reserves of prairie where scientific investigations could be made on problems concerning the native vegetations, floras, and faunas of the various topographic, chmatic, and prairie districts throughout Iowa. Therefore, they were meant to serve as a reference point by which future generations could compare the in- fluence of man on Iowa since settlement (Hayden 1946, Moyer 1953, Aikman 1959, Landers 1966). Kalsow Prairie, 65 ha (160 acres) of un- plowed grassland in Pocahontas County, Iowa, was purchased for these reasons in 1949. Since its purchase in 1949, it has been the object of studies on 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, Brotherson and Landers 1976, Brotherson 1980). 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 prairie vegetation. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 246 June 1982 Brotherson: Mima Mound Vegetation 247 Moyer (1953), Aikman and Thorne (1956), Ehrenreich (1957), Kennedy (1969), and Brotherson (1969) in more recent studies present ecological and taxonomic descrip- tions of four state-owned native prairie tracts. 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. 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 community. Steiger (1930) and Cain and Evans (1952) mapped the spatial distributions of several species. Tliey 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 tlie dispersal and establishment of new individuals. Local distribution patterns of species have been of interest to many ecologists. One factor influencing such distribution patterns in prairies across North America are mima mounds (Collins 1975, Del Moral 1976, and Mielke 1977). Such mounds, originally thought to be Indian burial mounds (Davids 1967), have been known to exist for many years. Their origin has frequently been con- tested in scientific literature. They have been considered the result of fossorial mammal ac- tivity, hydrostatic pressure, wind deposition, or several ground-frost phenomena (Scheffer 1947, Thorp 1949, Tester and Marshall 1961, Hansen 1962, and Davids 1967). Scheffer (1958), McCinnies (1960) and Ross et al. (1968), in reviews of mound development, in- dicate that none of the hypotheses con- cerning their origin is universally accepted. Ross et al. (1968) indicate that this dis- agreement is probably due to the description of different causes or combinations of causes at different locations. This study was undertaken to provide in- formation on the phytosociology of this par- ticular prairie phenomenon. It includes infor- mation on species composition and distribution of the mounds and on the mounds' relationship to the prairie vegetation. Methods This study was begun in the spring of 1967 and continued through the following year (1968) and into the summer of 1969. The study site (Kalsow Prairie) is one of four state-owned Iowa prairies. It is five miles northwest of Manson, Iowa, and comprises the NE 1/4 of Section 36, Belleville Township, T 90 N, R 32 W, Pocahontas County. It oc- curs in a part of north central Iowa that was glaciated during the most recent advances of the Wisconsin Glacier and within the Clari- on-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) and the pres- ence of mima mounds found scattered across the 65 ha of the study area. The vegetation of mima mounds was ana- lyzed using two separate approaches. The first involved the identification and listing of all species found within their boundaries. The second utilized random plots to determine percent cover, composition, and interspecific relationships of species within these subcommunities. Voucher specimens were collected in du- plicate throughout the growing seasons of 1967 and 1968. All specimens were identified and identical sets have been deposited in the herbaria of Iowa State University, Ames, Iowa, and Brigham Young University, Provo, Utah. Nomenclature follows Pohl (1966) for the grasses, Gilly (1946) for the sedges, and Gleason (1952) for the forbs. Quadrat Analysis The vegetation of each mound was sam- pled by using a 20 X 50 cm (1000 cm^) quad- rat (Fig. 3). The quadrats were located on a restricted basis to reduce bias and to keep ad- jacent quadrats at fairly equal distances 248 Great Basin Naturalist Vol. 42, No. 2 apart. Sampling was done between 1 August and 15 September each year when most spe- cies had reached their maximum growth. Cover estimates were made for each quadrat through use of Daubenmire'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 •WEST overlap of each species regardless of super- imposed canopies of other species. The can- opies of different species are commonly in- terlaced or superimposed over the same area; therefore, coverage percents often total greater than 100 percent. The mounds were first located (Fig. 1) and permanently identified by a numbered stake placed at the west edge of the mound. Every MAP OF TKE KALSOW PRAIRIE O MIMA MOUNDS • MIMA MOUNDS ADJACENT TO WHICH THE PRAIRIE WAS SAMPLED A AREAS AFFECTED BY SOIL DRIFT FROM ADJACENT FIELDS - - 20 ACRES OF PRAIRIE INVOLVED IN SOIL AND PLANT -^DISTRIBUTION STUDIES <9>* POTHOLES AND DRAINAGE Fig. 1. Map of Kalsow Prairie showing the distribution of mima mounds. June 1982 Brotherson: Mima Mound Vegetation 249 mound was then sampled starting at the northeast comer of the mound and gridding the mound with quadrats placed every three steps. The number of samples varied with the size of the mound, ranging from 5 on the smallest to 45 on the largest. A total of 1549 samples was taken on 128 mounds. Mound dimensions were taken in north-south and east-west directions, and areas (in square feet) were obtained through the use of the ellipse area fonnula A = pab A is the area; p = 3.1417; a is the length; and b is the width of the mound. Data was also taken to describe the prairie commimity adjacent to the mounds. This in- volved 444 samples taken adjacent to 37 se- lected miina mounds. Each mound was bi- sected by two transects oriented in north- south and east-west directions. Quadrats were then taken along these transects (Fig. 2) start- ing at the mound edge and progressing into the adjacent prairie. A total of 12 quadrats was taken adjacent to each mound, 3 in each direction. The quadrats were placed at 12- foot intervals. Data Analysis Data collected from quadrat studies, map- ping studies, and topographic studies were used to describe generally the vegetation of the mounds. Frequency values and average cover values were determined for all species in every stand. Frequency values were deter- mined by use of the following formula: Frequency (%) = Number of plots of occurrence Total number of plots sampled X 100 Cover values were determined by summing the midpoints of the cover-class ranges and dividing by the number of sample quadrats in the stand. An ordination technique proposed by Or- loci (1966) was employed to ordinate vegeta- tion units within the different sub- commimities listed above. Raw data were first summarized by hand calculation and then transferred to punch cards. This tech- nique was completed on an IBM S360 Mod 65 computer. Through this technique the en- tities to be ordinated (i.e., plant species or stands of vegetation) are projected as points into n-dimensional space. Such points are positioned by attribute scores through the ap- plication of the R- and Q-techniques of factor analysis. Once established, this multi- dimensional array of points is then reduced to a three-dimensional system. This is accom- plished by selecting 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 consid- eration are then positioned linearly in rela- tionship to these two extremes. This action thus establishes the X-axis. The above process is repeated unt 1 all points have been estab- lished 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. Expressions of interspecific association were attempted utilizing Cole's Index (1949). Step one in the computation of the index in- volves the accumulation of 2 X 2 contin- gency tables. Actual calculation of the index involves the formulas discussed by Brotherson (1980). Tests of statistical significance were performed by means of the Chi-square test. 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. Graphic representation of data obtained from topographic studies and from ordination analysis was drawn by the computer. 0 0 0 o o o MOUND o o o Fig. 2. Location of plots in upland prairie adjacent to selected mima mounds. 250 Great Basin Naturalist Vol. 42, No. 2 ■SOUTH FEET Fig. 3. Mlcrorelief, determined in July 1968, of Mound 14 (Fig. 1), a typical mima mound, Kalsow Prairie. Results Historical information as well as evidence obtained in this study indicates that much of the Kalsow Prairie has been subjected to mowing, grazing to some extent, and abun- dant pocket gopher activity. The distribution of mima mounds on the upland prairie is shown in Figure 1. The mima mounds, which are widely scattered across the 65 ha of Kal- sow Prairie, are numerous small circular mounds of soil ranging in diameter from 6 to 72 feet with a microrelief of from 6 to 36 inches (Fig. 3). They support a somewhat dif- ferent vegetation than the surrounding prairie. The mima mounds of Kalsow Prairie are low, rounded mounds of loose, nonstratified soil that occur most frequently on the higher, better-drained soils. Their origin is at present not well understood, but it seems that they are most likely initiated by activity of the pocket gopher (Geomys bursarius) and other animals. The activity of the pocket gopher on a selected 20-acre tract of the prairie is shown in Figure 4. Note the similarity in mima mound distribution on the same 20 acre unit (Fig. 5). Once initiated, the mounds might then be affected and enlarged by the differential expansion and contraction of their soils and by wind deposition of dust car- ried in from the adjacent cultivated fields. Continued use by pocket gophers and other burrowing mammals is evidenced by an abundance of recent soil pushed out from burrows in and about the mounds. A vegetation analysis of several mounds picked randomly as a representative sample of the mound phenomena on the Kalsow Prairie showed that the number of plant spe- cies supported by the mounds was only slightly greater than that of the surrounding prairie (i.e., mounds = 51 species, adjacent prairie = 49 species). Of these, 38 or 75 per- cent of the sampled species were found in common on mounds and prairie. Those spe- cies showing cover values greater than one are placed in Table 1. Indicator species were chosen as representative of the two areas and assigned adaptation numbers according to the method of Dix and Butler (1960). This infor- mation was then used to compute Plot Index Values (PIV) for the two areas and thus sepa- rate them spatially as shown in Figure 6. The Plot Index Values were computed by use of the following equation: ^^^ •• • •• •• ^ 1 • •• • •• ••• •••••• 1 • ••• • 1 ** > ••• • • • J ^^ » • •• ^ r^ — r •• • r^\ r ( ••• •••• » •• 1 \J — N •••••• • J \ •••• ••• • • ' /^ ■ — ^ \ ••• / / • \ ) •••• / / • • • •k^ / • • / / •••••• •••^~2, / •• y V. •.:::::::..C \ .:: :: >••••••• 1 :: .:::'::::;:\ 1 ••••••• >••••••• 1 • ••••• •• \ >••••• • • •••••••• \ ••• • • •• "-^ • •••••••• !•• ••••• .^i ^•* ••••• • •• • •• • • Fig. 4. Distribution of pocket gopher [Goemijs bur- sauziiis) activity in the 8 ha (20 acre) intensive study plot. T— ; y • • • f c • • • . f 1 p • J^ J • » J? • ]> •• •• { • • • • • • • •• • •• • •• { >• • • •• v \ • •• • ♦ • • > V ♦ • • \ T • \j • • \ -^^= ^-^ •• Fig. 5. Distribution of mima mound influences in the ha (20 acre) intensive study plot. June 1982 Brotherson: Mima Mound Vegetation 251 Sum (percent cover of each indicator species X its adaptation # ) PIV = — Sum (percent cover of each indicator species) This spatial separation (Fig. 6) and Table 1 indicate definite differences in the vegetation of the two areas. To further strengthen the hypothesis that the vegetation changes from the mounds to the prairie, a similarity index was computed for the two areas utilizing So- rensen's (1948) index of similarity. The value of K was calculated to be 35.2 percent, which means that the mound vege- tation and the adjacent prairie vegetation have a similarity of 35 percent. Similarity values reported by Curtis (1959) for a series of communities in Wisconsin showed ex- tremes from 34.1 to 70.3. It seems, therefore, that mound vegetation is quite distinct from that of the prairie. The relationship of mound vegetation to adjacent prairie has not been extensively in- vestigated (Ross et al. 1968). Attempts to de- scribe such relationships in the present study revealed that the mounds are associated with changes in the surroimding prairie vegeta- tion. These changes were investigated by sampling prairie vegetation adjacent to 37 of Table 1. Average percentage cover values in mound and adjacent prairie areas for all species with a percent cover greater than one. % Cover % Cover Species (prairie) (mounds) 1 2 Sporobohis heterolepis Amorpha canescens Schizachijrium scoparius Zizia aurea 53.60 2.00 1.00 1.20 .04 .83 .83 Aster laevis 1.10 .04 Solidago rigida 1.30 .10 2 3 Paniciim leihergii Aster ericoides 3.00 3.00 1.80 4.00 3 Andropogon gerardi Ratibida colwnnifera 14.00 1.00 11.85 2.90 Achillea millifolium .93 2.40 4 Phy sails heterophylla Rosa stiff ulta .04 .80 1.90 1.70 4 Convolvidiis sepium .10 1.90 5 Asclepias syriaca Agropyron repens Ambrosia artemisifolia Solidago canadensis 3.00 1.10 1.40 4.30 8.36 5 Poa pratensis 8.00 45.26 *These numbers are the adaptation numbers assigned to the different in- dicator species. the 128 mounds studied. Two transects, one oriented north-south and the other east-west, were extended through each mound. The ad- jacent prairie vegetation was sampled along these transects starting at the mound bound- ary and extending into the prairie. Twelve samples at 12-foot intervals were taken adja- cent to each mound, as shown in Figure 2. Cover estimates were recorded for each spe- cies present in the quadrat. The resulting data were analyzed by grouping all quadrats found at equal intervals from the mounds and averaging them to obtain percentage cover values for all participating species (Table 2). A similar analysis was also completed by grouping all quadrats located on the north, south, east, and west sides of the mounds and again averaging to obtain percentage cover values for all participating species (Table 3). In both cases it is evident that the mounds have provided a new microenvironment to which some prairie species respond. Some species (Table 2) show a positive response by appearing almost exclusively on the mounds or by increasing in importance from the prairie toward the mound. Species showing this type of response were Achillea millifo- lium, Agropyron repens. Ambrosia artemisi- folia, Asclepias syrica. Aster ericoides, Cheno- podium alburn. Convolvulus sepium, Elymus canadensis, Fragaria virginiana, Galium oh tusum, Helianthus grosseserratus, Helianthus laetiflorus, Heliopsis helianthoides, Oxalis stricta, Physalis heterophylla, Poa pratensis, Ratibida columnifera, Rosa suffidta, Solidago canadensis, and Spartina pectinata. Other species showed a negative response decreas- ing in importance as the mound is ap- proached from the prairie. These species were Amorpha canescens. Aster laevis, Rap- tisia leucantha, Comandra unibellata, Eryn- gium yuccifolium, Lathyrus venosus, Liatris pycnostachya, Silphium laciniatum, Solidago rigida, Sporobolus heterolepis, and Zizia aurea. Another response is exhibited by An- dropogon gerardi. It increases in importance as you move toward the mound then drops Fig. 6. Linear ordination of the mima mound and ad- jacent prairie vegetation according to their plot index values. A = mima mounds, B = adjacent prairie. 252 Great Basin Naturalist Vol. 42, No. 2 Table 2. Effects of mima mounds on cover values of the surrounding upland prairie vegetation. Distance from mound Species )und 12 ft 24 ft 36 ft .74 .50 .59 .51 1.16 .02 .02 .03 .01 .00 .00 .00 1.91 .02 .02 .00 .41 .00 .00 .00 .41 1.94 2.01 1.35 .14 .02 .00 .00 7.69 18.72 14.66 12.80 .00 .00 .00 .02 .01 .10 .06 .02 .07 .10 .00 .12 .01 .00 .00 .00 .61 .49 .30 .25 .01 .00 .00 .00 1.18 .17 .02 .00 .35 .51 .19 .03 .04 .03 .02 .00 3.96 2.23 2.50 1.82 .97 1.60 1.76 .81 .51 .30 .20 .24 .00 .02 .02 .00 .05 .10 .17 .02 lOO .00 .10 .10 .03 .07 .05 .02 .00 .00 .00 .10 .01 .00 .00 .02 .19 .10 .07 .16 .00 .02 .00 .00 .00 .00 .00 .02 .77 .02 .00 .00 .00 .00 .03 .00 .50 .03 .19 .15 .25 .00 .00 .00 .06 .23 .27 .34 3.13 .53 .28 .22 1.25 2.21 1.91 1.79 .00 .00 .00 .02 1.22 .42 .47 .39 .06 .00 .00 .00 .07 .07 .14 .07 .01 .00 .00 .00 .01 .10 .14 .02 .01 .00 .00 .00 .83 .79 .63 .41 .93 .74 .69 .54 .00 .02 .05 .15 4.13 1.72 1.67 2.36 1.59 1.52 1.23 1.03 .46 .14 .02 .00 1.32 .51 .37 .63 .00 .02 .00 .00 .13 .00 .00 .00 .12 .00 .00 .02 .02 .00 .03 .02 .13 .20 .19 .07 .00 .02 .04 .04 .06 .02 .04 .04 .01 .20 .19 .29 Achillea millifolium Agropyron repens Agropyron traclnjcaubim Ambrosia arternisifolia Ambrosia trifida Amorpha canescens Amphicarpa bracteata Amlropogon gerardi Anemone canadensis Anemone cylindrica Apocynum sibiricum Arabis hirsuta Artemisia ludoviciana Asclepias sidlivantii Asclepias syriaca Asclepias tuberosa Asclepias verticillata Aster ericoides Aster laevis Aster simplex Astragalus canadensis Baptisia leucantha Baptisia leucophaea Boutelotia curtipendula Caenothus americanus Calamagrostis canadensis Carex gravida Carex aquatilis Carex retrorsa Chenopodium album Ciciita maculata Cirsium altissimum Cirsium arvense Comandra umbellata Convolvulus sepium Desmodium canadense Echinacea pallida Elymus canadensis Elymtis virginicus Equisetum kansanum Erigeron strigosus Eryngium yuccifolium Euphorbia serpyllfolia Fragaria virginiana Galium obtusum Gentiana andrewsii Helianthus grosseserratus Helianthus laetiflorus Helianthus maximiliani Heliopsis helianthoides ] uncus tenuis Kochia scoparia Lactuca scariola Lathyrus palustris Lathy rus venosus Lespedeza capitata Liatris aspera Liatris pycnostachya June 1982 Table 2 coninued. Brotherson: Mima Mound Vegetation 253 Species Distance from mound lound 12 ft 24 ft 36 ft .24 .57 .54 .37 .08 .03 .02 .03 .00 .10 .02 .10 .00 .00 .02 .02 .40 .51 .69 .39 .11 .00 .00 .00 1.42 .08 .02 .02 .27 .00 .00 .00 2.02 3.13 3.89 2.52 .62 .52 .39 .54 .00 .00 .02 .03 .02 .19 .10 .25 .02 .25 .35 .20 .01 .00 .03 .03 .19 .25 .41 .20 1.24 .12 .08 .02 .35 .10 .14 .07 40.47 10.14 7.64 6.69 .14 .00 .00 .00 .09 .00 .00 .00 .09 .00 .10 .02 .15 .14 .24 .10 .00 .07 .00 .19 3.08 2.19 1.30 1.49 2.48 1.20 1.49 1.50 .01 1.37 .42 .46 .03 .03 .04 .02 .13 .03 .03 .14 .62 .00 .02 .00 .03 .00 .00 .00 .02 .39 .54 .69 .09 .00 .00 .00 10.73 5.03 3.80 4.51 .00 .02 .00 .02 .11 .23 .35 .20 .47 1.37 2.50 1.47 .00 .10 .27 .65 .94 .37 .32 .24 1.05 22.61 31.00 33.97 .25 .59 .89 .49 .00 .10 .00 .00 .01 .00 .00 .00 .01 .00 .00 .00 .05 .00 .02 .00 .19 .27 .19 .27 .13 .07 .10 .05 .11 .30 .07 .14 1.44 2.08 2.47 1.65 Lithospennum canescens Lysimachia chiliata L ijsimach ia hyb rida Lysimachia qitadriflora Muhlenbergia racemosa Oenothera biennis Oxalis stricta Panicum capillare Panicum leibergii Panicum virgatum Pedicukiris canadensis Petalostemum candidum Peta lostem am piirpureiim Phleum pratensis Phlox pilosa Physalis heterophylla Phijsalis virginiana Poa pratensis Pohjgoniim cotnolvuhts Pohjgonum ramosissimiim Potentilla arguta Psoralea argophylla Pycnanthemiim virgiiiianum Ratibida cohimnifera Rosa suffidta Schizachyrium scoparium Scutellaria leonardii Senecio pauperculus Seturia lutescens Setaria viridis Silphium laciniatum Solanum nigrum Solidago canadensis Solidago gy mnospennoides Solidago missouriensis Solidago rigida Sorghastrum nutans Spartina pectinata Sporobolus heterolepis Stipa spartea Taraxacum officinale Thalictrum dasycarpum Tradescantia bracteata Trifolium pratense Viola pedatifida Viola sp. Vicia americana Zizia aurea sharply in average percentage cover as you reach the mound proper. Other species show- ing this same kind of response were Schiz- achyriiwi scoparius, Bouteloiia curtipendula, Desmodiurii canadense, and Lithospermum canescens. Evidence from Table 3 indicates that sev- eral species also showed some response to small differences in microrelief as associated with aspect. Species showing preference for the southern aspect were Amorpha canes- cens, Schizachyrium scoparius, Asclepias tuberosa. Aster ericoides, Comandra umbel- lata, Lithospermum canescens, Panicum lei- bergii, Petalostemum candidum, Poa pra- tensis, Solidago missouriensis, and Stipa 254 Great Basin Naturalist Vol. 42, No. 2 Table 3. Effects of iniina mounds on cover values of the surrounding upland prairie vegetation in relation to aspect. Species Mound North South East West Achillea millifolium Agropyron repens Ambrosia artemisifolia Amorpha canescens Amphicarpa bracteata Andropogon gerardi Anemone canadensis Anemone cylindrica Apocynum sibiricum Artemisia ludoviciana Asclepias syriaca Asclepias tuberosa Asclepias verticillata Aster ericoides Aster laevis Aster simplex Astragalus canadensis Baptisia leucantha Bouteloua curtipendiiki Calamagrostis canadensis Carex brevoir Carex aquatilix Carex lasiocarpa Chenopodium album Cicuta maculata Cirsium altissimum Comandra umbellata Convolvulus sepium Desmodium canadense Echinacea pallida Elymus canadensis Equisetum kansanum Eryngium yuccifolium Fragaria virginiana Galium obtusum Gentiana andrewsii Helianthiis grosseserratus Helianthiis laetiflorus Helianthus maximiliani Heliopsis helianthoides Juncus tenuis Lactuca scariola Lathyrus palustris Lathyrus venosus Lespedeza capitata Liatris aspera Liatris pycnostachya Lithospermum canescens Lysimachia chiliata Lysimachia hybrida Lysimachia quadriflora Muhlenbergia racemosa Oxalis stricta Panicum leibergii Panicum virgatum Pedicularis canadensis Petalostemum candidwn .74 .56 .38 .68 .56 1.16 .02 .00 .05 .02 1.91 .02 .02 .00 .00 .41 2.16 2.68 1.24 2.07 .41 .00 .00 .02 .00 7.69 16.24 15.16 16.64 13.22 .00 .00 .00 .02 .00 .01 .11 .05 .07 .05 .07 .11 .00 .07 .00 .61 .36 .25 .45 .34 1.18 .16 .05 .00 .02 .35 .14 .74 .02 .11 .04 .02 .00 .02 .02 3.96 2.09 2.52 2.00 2.17 .97 1.31 1.31 1.62 1.55 .51 .14 .11 .27 .47 .00 .05 .00 .00 .00 .05 .00 .00 .29 .14 .03 .02 .02 .09 .05 .01 .11 .00 .02 .00 .19 .00 .11 .23 .05 .00 .00 .02 .00 .00 .00 .00 .00 .02 .00 .77 .00 .00 .00 .02 .00 .00 .00 .00 .05 .50 .09 .07 .07 .20 .06 .20 .50 .45 .23 3.13 .63 .25 .23 .47 1.25 1.78 1.91 1.89 2.16 .00 .00 .02 .02 .00 1.22 .43 .63 .45 .25 .07 .11 .11 .09 .07 .01 .05 .00 .14 .14 .83 .59 .70 .68 .65 .93 .88 .86 .52 .68 .00 .05 .14 .02 .05 4.13 2.41 2.18 2.09 1.01 1.59 .97 1.26 1.58 1.22 .46 .16 .00 .05 .02 1..32 .43 .52 .92 .27 .00 .00 .00 .02 .00 .12 .02 .00 .00 .00 .02 .02 .00 .05 .02 .13 .07 .14 .23 .05 .00 .07 .00 .07 .02 .06 .00 .02 .09 .05 .01 .16 .29 .23 .27 .24 .43 .74 .41 .45 .08 .00 .05 .05 .02 .00 .00 .16 .02 .00 .00 .00 .00 .02 .02 .40 .56 .47 .36 .79 1.42 .02 .02 .07 .05 2.02 2.97 4.35 2.57 2.68 .62 .36 .63 .65 .41 .00 .00 .00 .05 .05 .02 .18 .27 .14 .11 June 1982 Brotherson: Mima Mound Vegetation 255 Table 3 continued. Species Mound North South East West Petalosteinwn purpureum Phletim pratense Phlox pilosa Physalis hcterophijUa Physalis virginiana Poa pratensis Potentilla arguta Psoralea argophylla Pycnanthemian virginianum Ratibida cohannifera Rosa sitffulta Schizachyriiim scopariiim Scutellaria leonardii Senecio pauperculus Setaria lutescens Silphium laciniatum Solidago canadensis Solidago gymnospennoides Solidago missouriensis Solidago rigida Sorglvstnan nutans Spartina pectinata Sporoboliis heterolepis Stipa spartea Taraxacum officinale Trifolium pratense Viola pedatifida Viola sp. Vicia americana Zizia aurea .02 .14 .16 .52 .29 .01 .05 .02 .02 .02 .19 .34 .18 .45 .20 1.24 .00 .14 .11 .23 .35 .14 .07 .16 .14 40.47 7.55 9.89 7.76 6.42 .09 .02 .02 .14 .00 .15 .09 .20 .32 .09 .00 .05 .05 .05 .07 3.08 1..37 1.80 2.00 1.60 2.48 1.55 1.55 .83 1.22 .01 .45 1.28 .86 .45 .03 .05 .02 .07 .02 .13 .18 .02 .07 .05 .62 .00 .02 .00 .00 .02 .29 .61 .36 .90 10.73 5.50 4.28 4.12 4.08 .00 .00 .00 .07 .00 .11 .20 .52 .27 .11 .47 1.94 1.64 1.46 2.03 .00 .95 .14 .16 .27 .94 .14 .45 .56 .25 1.05 32.30 27.03 24.28 36.03 .25 .18 .50 ..38 .38 .00 .14 .00 .00 .00 .05 .00 .00 .02 .00 .19 .27 .27 .29 .16 .13 .07 .09 .02 .14 .11 ..36 .18 .09 .14 1.44 2.45 2.05 2.30 2.36 spartea. Species showing preference for the north side of the mounds were Anemone cy- lindrica, Asclepias syriaca. Convolvulus se- pium, Sorghastrum nutans, and Vicia americana. It seems the mounds, however sHght in mi- crorehef, provide sufficient modification of the prairie to allow striking patterns of vege- tational change to emerge. In one aspect the mounds provide habitats that exhibit differ- ent levels of disturbance (i.e., the amount of actual disturbance decreases as you leave the mound and proceed into the prairie). The re- sponse of several species to the disturbance factor would tend to support the hypothesis that in the Kalsow prairie the changes that have occurred since 1953 are in effect caused by some degree of disturbance. Several of the species that showed increased importance since Moyer's (1953) work (i.e., Solidago canadensis, Panicum leibergii, Helianthus grosseserratus, Desmodium canadense, Ga- lium obtusum, and Fragari,a virginiana) also showed a corresponding increase in impor- tance as you approach the mounds from the prairie. Likewise, some of the species which decreased in importance in the past 16 years (i.e., Zizia aurea, Sorghastrum nutans, and Sporobolus heterolepis) decreased as distur- bance increased. Several species, Andropogon gerardi, Schizachyriwn scoparius, Bouteloua curtipendula, and Lithospennum canescens, indicated positive response to slight disturb- ance but negative response to heavier disturbance. It seems, therefore, that mounds or other forms of disturbance affect vegetation changes in the prairie which, when consid- ered over a period of years, may alter the original structure and composition of its veg- etation. Whether such changes would be per- manent or temporary is a question that can be answered only by long-term studies set up to follow the fluctuations of the prairie vege- tation and its environment. 256 Great Basin Naturalist Vol. 42, No. 2 Fig. 7. Three-dimensional ordination of 128 mima mounds found in Kalsow Prairie. Analysis of the mound vegetation as a unit was attempted using Orloci's (1966) method of ordination. Each mound was considered as a stand of vegetation and all 128 mounds were projected into three-dimensional space (Fig. 7). This analysis placed the 128 mounds into a relatively linear relationship in the X, Y, and Z planes (Fig. 7). This indicated that only two to three factors could be responsible for the placing of each mound into this sort of an alignment in relation to all the other mounds. Further study indicated that align- ment was closely related to the two species Poa pratensis and Solidago canadensis. Poa pratensis was responsible for alignment of the X-axis (Fig. 8) and Solidago canadensis was responsible for alignment of the Y-axis (Fig. 9). Because no environmental measurements were taken, it was impossible to determine the causative factors to which these two spe- cies were linked. It appears, however, that the vegetation of the mounds fits the concept of a continuum and that perhaps the con- trolling environmental factors would be re- lated to the age of the mound and the degree of disturbance. Poa pratensis, for example, is an in- troduced species whose characteristics are such that it is able to compete well within the environment of the prairie protected from early spring fires. Under conditions of grazing, mowing, and other disturbance, Poa pratensis is known to increase in importance (Weaver 1954). By ordinating the species of the mounds into three-dimensional space (Fig. 10), it was found that only those species having irregular distribution patterns were isolated. The most important species were Poa pratensis, Soli- dago canadensis, Solanum nigrum, Andropo- gon gerardi. Aster ericoides, Helianthus grosseserratus, Connvolvulus sepium, He- lianthus laetiflorus, Desmodium canadense. Ambrosia artemisi folia, Panicum leibergii, Ratibida columnifera, Rosa suffulta, and Zizia aurea. Again it was found that the two species Poa pratensis and Solidago canadensis were responsible for alignment of the X and Y-axes. To further understand the relationships of mound vegetation, interspecific association values were computed for all possible pairs of species (Table 4). Out of 7200 possible com- binations only 78, or about 1 percent, showed any degree of positive association. Basic clus- ters or groups of species within these 78 posi- tive association units are illustrated in Fig- ures 11 and 12. Fig. 8. Two-dimensional ordination of mima mounds with percentage cover values of Poa pratensis shown re- lating indirectly to the X-axis. Fig. 9. Two-dimensional ordination of mima mounds with percentage cover values of Solidago canadensis shown relating indirectly to the Y-axis. June 1982 Brotherson: Mima Mound Vegetation 257 Five of the six clusters (B through F) shown in Figure 1 1 are composed of species that are generally tolerant of disturbance and were shown to be most important on the mound it- self (Table 2). Cluster A (Fig. 11) is composed of upland prairie species. Figure 12 shows four clusters that are designated as A, B, C, and D. Cluster A has as its center Solidago canadensis and as associated species Carex gravida, Monarda fistulosa, Senecio pauper- culus. Aster simplex, Solanum nigrum, and Apocynum sibiricum. Cluster B has for its center a unit of three species: Desmodium canadense, Fragaria virginiana, and He- lianthus grosseserratus. These are then associ- ated with Galium obtusum and several other species only on a very limited basis (i.e., 20 to 40 percent). Cluster C has as its center Zizia aurea and as associated species: Lythrum ala- tum, Pedicularis canadensis, and Petaloste- mum, candidum. Here again Zizia aurea and its associated species are weakly associated with several other species as well as to clus- ter D. Cluster D is basically a discrete unit including Lycopus americanus, Lysimachia chiliata, and Spartina pectinata. In all cases the clusters of Figure 12 appear to be com- posed of species generally found on lowland prairie soils or bordering potholes and drain- age ways. This would indicate that the mound environment as shown by these groups might be somewhat more moist than the adjacent prairie. Specifically, the interspecific association analyses indicate two groups of species oc- cupying the mima mounds of Kalsow prairie. 100-- 50-- o" A ®i<« '- btfo. ef""o' S- ^ tr 50 100 530 Y Fig. 10. Two-dimensional ordination of species found in the mima mound study; A = cluster of species not show- ing distinct distribution patterns, b = Solanum nigrum, c = Ambrosia trifida, d = Agropyron repens, e = Amorpha canescens, i = Chenopodium album, g = Fragaria virginiana, h = Achillea millifolium, i = Asclepias syriaca, j = Spartina pectinata, k = Panicum virgatum, 1 = Zizia aurea, m = Heliopsis helianthoides, n = Physalis heterophylla, o = Elymus canadensis, p = Aster laevis, q = Oxalis stricta, r = Artemisia ludoviciana, s = Sporobolus heterolepis, t = Aster simplex, u = Galium obtusum, v = Ambrosia artemisifolia, w = Convolvulus sepiurn, x = Helianthus lae- tiflorus, y = Desmodium canadense, z = Ratibida columnifera, aa = Panicum leibergii, bb = Rosa suffulta, cc = Helianthus grosseserratus, dd = Aster ericoides, ee = Andropogon gerardi, ff = Solidago canadensis, gg = Poa pratensis. 258 Great Basin Naturalist Voh-42, No. 2 Table 4. Coles Index values expressing positive interspecific association in niima mound communities. Species Species X2a Ct^ Gv- Agropyron repens Amorpha canescens Andropogon gerardi Apocynitm sibiricum Artemisia hidoviciana Asclepias tuberosa Aster laevis Aster simplex Bouteloiia curtipendula Carex gravida Chenopodium album Cirsium altissimum Comandra iimbellata Desmodium canadense Equisetitm kansanum Erijngiwn yuccifolium Fragaria virginiana Galium obtusum Helianthiis maximiliani Kochia scoparia Lactuca scariola Lathy riis pahistris Liatris pycnostachya Lithospermum canescens Lycopus americanus Convolvulus sepium Panicum leibergii Sporobolus heterolepis Poa pratensis Solidago canadensis Spartina pectinata Convolvulus sepium Panicum virgaturn Panicum leibergii Elymus canadensis Hclianthus grosseserratus Ratibida cohimnifera Solidago canadensis Convolvulus sepium Lithospermum canescens Panicum leibergii Phlox pilosa Zizia aurea Fragaria virginiana Physalis virginiana Solidago canadensis Elymus canadensis Panicum cap ilia re Desmodium canadense Fragaria virginiana Calium obtusum Panicum leibergii Sporobolus heterolepis Fragaria virginiana Galium obtusum Helianthiis grosseserratus Panicum virgaturn Ratibida cohimnifera Solidago canadensis Spartina pectinata Zizia aurea Helianthiis laetifloriis Viola pedatifida Galium obtusum Helianthiis grosseserratus Helianthiis grosseserratus Panicum virgatum Zizia aurea Panicum leibergii Physalis heterophylla Phlox pilosa Ratibida cohimnifera Solidago rigida Lithosperiniim canescens Petalostemurn purpureiim Psoralea argophylla Solidago rigida Sporobolus heterolepis Panicum leibergii Lysiinachia chiliata Spartina pectinata *Chi-square "Cole's Index •^Standard deviation Cole's Index 48.20 .45 .06 37.20 .49 .07 24.82 .19 .03 8.82 .63 .21 5.04 .33 .14 15.77 .21 .05 16.97 .19 .04 12.68 .17 .04 23.91 .19 .03 5.99 .19 .07 54.44 .35 .04 8.92 .17 .05 17.50 .45 .11 3.80 .36 .18 14.34 .23 .06 4.13 .30 .14 9.26 .21 .07 4.10 .21 .10 3.91 .19 .09 17.76 .25 .05 7.54 .65 .23 7.69 .18 .06 39.39 .17 .02 14.88 .20 .05 17.87 .26 .06 22.35 .29 .06 10.20 .32 .09 28.49 .25 .04 230.53 .45 .02 239.96 .46 .02 75.32 .29 .03 55.13 .17 .02 20.91 .19 .04 8.52 .22 .07 79.85 .24 .02 87.88 .31 .03 13.69 .34 .09 75.09 .82 .09 137.95 .30 .02 71.48 .24 .02 72.33 .24 .02 75.68 .17 .01 68.88 .24 .02 4.31 .23 .11 9.38 .42 .13 5.36 .17 .07 5.58 .36 .15 5.68 .19 .08 19.50 .36 .08 23.57 .19 .03 11.75 .18 .05 7.77 .25 .08 6.69 .24 .09 16.28 .25 .06 145.04 .66 .05 9.46 .62 .20 June 1982 Brotherson: Mima Mound Vegetation 259 Table 4. Cole's Index values expressing positive interspecific association in minia mound communities. Species Lijsimachia chiliata Lijthnim alatitm Monarda fistitlosa Oenothera biennis Paniettm viroatuni Pediciilaris canadensis Petalostemtim eandiduni Polygonum raniosissimum Potentilki arguta Senecio paupercuhis Silphium laciniatum Solanian nigrum Solidago rigida Viola sp. Species Spartina pectinata Zizia aurea Zizia aurea Solidago canadensis Panicum capillare Solidago canadensis Spartina pectinata Zizia aurea Zizia aurea Rumex crispus Viola pedatifida Solidago canadensis Solidago rigida Spartinia pectinata Zizia aurea Solidago rigida Spartina pectinata Viola sp. Zizia aurea Solidago canadensis Zizia aurea Zizia aurea X2a C,b Gt^ 40.75 .60 .09 12.21 .40 .11 20.90 .70 .15 5.17 1.00 .43 102.87 .33 .03 20.30 ..37 .08 27.11 .15 .02 26.42 1.00 .19 10.36 .30 .09 71.91 .22 .02 7.62 .17 .06 6.56 .70 .27 4.90 .17 .07 10.20 .31 .09 8.91 ..36 .12 7.60 .19 .06 11.51 .30 .08 97.96 .42 .04 9.31 .33 .10 4.29 .51 .24 23.58 .19 .03 17.37 .30 .06 ^Chi-square Cole's Index '^Standard deviation Cole's Index Fig. 11. Association of species found in the mima mound study, Kalsow Prairie, as determined by Cole's (1949) Index, the more lines between species, the greater the association; (A) Er yu = Enjngium yuccifoliitm, Vi pe = Viola pedatifida, Po ar = Potentilla arguta, (B) Ko sc = Kochia scoparia, Ph he = Physalis heterophylla, (C) Eq ka = Equisetutn kansanum. He la = Helianthus laeti floras, (D) An ge = Andropogon gerardi, Po pr = Poa pratensis, (E) Ci al = Circium altissimum, Pa ca = Panicum capillare, Oe bi = Oenothera biennis, (F) Po ra = Polygonum ramosissimum, Ru cr = Rumex crispus. Those that are tolerant to disturbance and are found occupying the central portions of the mound proper (Fig. 11) and those that are characteristic of lowland prairie areas and are more important at the edges and lower slopes of the mounds. We could find no indication that these groups were associ- ated with mound size. It appears that once a mound is formed, a new microenvironment is created that affects directly the structure and stability of the sur- rounding prairie. This effect is shown in the response of many species to the creation of these new habitats. The factors deemed most influential in affecting these new habitats are disturbance and microrelief. Field observa- tions indicate that the mounds represent mi- crosuccession sites and cause changes in the prairie vegetation to earlier stages in the sere. This hypothesis is supported by the fact that, in all cases studied, the vegetation of the mounds included a number of weed spe- cies (annuals, biennials, and some perennials) that are recognized as pioneer species. The resulting mound vegetation appears to be made up of a mixture of these pioneer species and species from the prairie that respond fa- vorably to mound disturbance. Present evi- dence also indicates that mound vegetation is 260 Great Basin Naturalist Vol. 42, No. 2 Fig. 12. Association of species found in the mima mound study, Kalsow Prairie, as determined by Cole's (1949) In- dex, the more Hnes between species, the greater the association; groups A, B, C, and D are basic clusters; Ag re = Agropijron repens. Am ca = Amorpha canescens, Ap si = Apocijnum sibiricum, Ar lu = Arteriiisia htdoviciana. As si = Aster simplex. As tu = Asclepias tuberosa. Bo cu = Boiitehua curtipendula, Ca ca = Calamagwstis canadensis, Ca gr = Carex gravida, Ch al = Chenopodium album, Co se = Convolvulus sepium, De ca = Desmodium cana- dense. El ca = Elymus canadensis, Fr vi = Fragaria virginiana, Ga ob = Galium obtusum. He gr = Helianthus grosseserratus. Mo fi = Monarda fistulosa. Pa le = Panicum leibergii. Pa vi = Panicum virgattim, Pe ca = Pedicu- laris canadensis. Pet ca = Petahstemum candidum, Ph Pi = Phlox pilosa, Ps ar = Psoralea argophijlla. La pa = Latlnjrus palustris. La sc = Lactuca scariola, Li ca = Lithospermum canescens, Li py = Liatris pijcnostachija, Ly al = Lijthrum alatum, Ly am = Lycopus americanus, Ly ch = Lysimachia chiliata, Ra co = Ratibida columnifera, Se pa = Senecio pauperculus. Si la = Silphium laciniatum, Sp he = Sporobolus heterophylla, Sp pe = Spartina pecti- nata. So ni = Solarium nigrum. So ca = Solidago canadensis. So ri = Solidago rigida, Vi sp = Viola sp., Zi au = Zizia a urea. undergoing succession that may be repeat- edly set back by more disturbance. It seems that mound vegetation is strongly influenced in its basic composition by the adjacent prairie flora. Literature Cited AiKMAN, J. M. 1959. Prairie research in Iowa. American 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, of Sci. 63:177-200. Brennan, K. M. 1969. Vertebrate fauna of Kalsow Prairie. Unpublished thesis. Iowa State Univ., Ames, Iowa. Brotherson, J. D. 1969. Species composition, distribu- tion, and phytosociology of Kalsow Prairie, a mesic tall-grass prairie in Iowa. Unpublished dis- sertation. Iowa State Univ., Ames, Iowa. 196 pp. 1980. Zonation patterns in the potholes of Kalsow Prairie, Iowa. Great Basin Nat. 40:327-384. Brotherson, J. D., and R. Q. Landers. 1976. 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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., Ames, Iowa. 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., Ames, Iowa. 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, Pennsylvania, Lan- caster Press, Inc. 3 vol. Hansen, R. M. 1962. Movements and survival of Tlw- jiKtmys talpoides in a mima mound habitat. Ecol- ogy 43:151-154. 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. Sci. 53:45-82. Kennedy, R. K. 1969. An analysis of tall-grass prairie vegetation relative to slope position, Sheeder Prairie, Iowa. Unpublished thesis. Iowa State Univ., Ames, Iowa. Landers, R. W. 1966. Visit the virgin prairie. Iowa Farm Science 21:418-419. McGinnies, W. J. 1960. Effect of mima type microrelief on herbage production of five seeded grasses in western Colorado. J. Range Manag. 13:231-239. MiELKE, H. W. 1977. Mound building by pocket gophers (Geomydoe): their impact on soils and vegetation in North America. J. Biogeography 4:171-180. Moyer, J. F. 1953. Ecology of native prairie in Iowa. Unpublished dissertation. Iowa State Univ., Ames, Iowa. Norton, D. C, and P. E. Ponchillia. 1968. Stylet- bearing nematodes associated with plants in Iowa prairies. J. Iowa Acad. Sci. 75:32-35. Orloci, L. 1966. Geometric models in ecology. I. 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Shimek, B. 1911. The prairies. Bull. Lab. Nat. Hist., State Univ. Iowa 6:169-240. 1915. The plant geography of the Lake Okoboji region. Bull. Lab. Nat. Hist., State Univ. of Iowa 7:1-90. 1925. The persistence of the prairie. Univ. of Iowa Studies 11:3-24. SoRENSEN, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on sim- ilarity of species content. Biol. Rev. 23:411-488. Steiger, T. L. 1930. Structure of prairie vegetation. Ecology 11:170-217. Tester, J. R., and W. H. Marshall. 1961. A study of certain plant and animal interrelations on a na- tive prairie in northwestern Minnesota. Min- nesota Museum of Natural History. Occasional Papers No. 8:1-50. Thorp, J. 1949. Effects of certain animals that live in the soil. Sci. Monthly 68:180-191. Weaver, J. E. 1930. Underground plant development in its relation to grazing. Ecology 11:543-557. 1954. North American Prairie. Johnsen Pub- lishing Company, Lincoln, Nebraska. 348 pp. 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. 1934. The prairie. Ecol. Monogr. 4:109-295. OCCURRENCE AND EFFECT OF CHRYSOMYXA PIROLATA CONE RUST ON PICEA PUNGENS IN UTAH David L. Nelson' and Richard G. KrebilF Abstract.- In a rare 1969 epidemic, spruce cone rust caused by Chrysomyxa piwlata infected 40-100 percent of trees and 20-67 percent of cones on riparian Colorado blue spruce on plots located in a 2200-2400 m elevational zone in Huntington Canyon of central Utah. Uredinial and telial sporulation on Ptjrola spp. began in mid-June, a time closely correlated with opening of pistillate spruce cones. Cone phenology and host habitat, as influenced by elevation, are apparently important factors in the restricted niche of the cone rust fungus in Utah. Several preceding consecutive years with extended periods of spring and fall moisture were associated with occurrence of the epidem- ic, although no cause-and-effect relationship was established. Weather records indicate that these events are in- frequent in this climatic zone, and there was no detectable recurrence of cone rust for at least 9 years following 1969. Outwardly normal seeds developed in diseased cones, but seed germinability was reduced by 25 percent. Ae- cial spore masses between cone scales, cone resinosis, and distortion of cone scales prevented seed dispersal to the extent that the seed crop was effectually destroyed. Our discovery of a rare outbreak of spruce cone rust caused by Chrysomyxa pirolata (Komike) Wint. on Picea pungens Engelm. in central Utah (Nelson and Krebill 1970) af- forded the opportunity to study the nature of the disease. Mycological collection records provide insight into its distribution; but infor- mation on the effect, ecological nature, and epidemiology of the disease is limited, espe- cially for the contiguous western United States. Review The spruce cone rust fungus is hetero- ecious and full cycled (Eraser 1912, Savile 1953, Ziller 1974). The aecial and pycnial stages form on female cones of Picea spp. Peridermial aecia develop on outer surfaces of cone scales (Arthur and Kern 1906). Myce- lium of the uredinial and telial stages is sys- temic and perennial in Pyrola and Moneses spp. (Rice 1927, Gaumann 1959). The per- ennial nature ensures persistence of the rust during periods unfavorable for infection (Sav- ile 1953). Favorable habitat for C. pirolata occurs primarily in the boreal regions of the north- ern hemisphere (Jcirstad 1940, Savile 1950, Ziller 1974), extending across North America and Eurasia. In the contiguous western United States, incidence is relatively low, and on spruce considered only occasional (Hedg- cock 1912) compared with Canada, where its occurrence is frequent (Can. Dept. Environ. 1951-1975). Abundance similar to that in Canada is evident in Alaska (Cash 1953, Kim- mey and Stevenson 1957, Zasada and Gre- gory 1969), and it occurs on Pyrola (pyrola) or Moneses (single-delight) species in Nevada (Arthur 1907-1931), California, Colorado, Idaho, Oregon, Montana, New Mexico, Utah, Washington, Wyoming (Arthur 1934), South Dakota (Peterson 1961), and Arizona (Gil- bertson and McHenry 1969). As in other re- gions of its distribution (Ziller 1974), in the western United States in certain areas, it is rather common on Pyrola and Moneses spp., but not known on spruce. Prior to this ac- count, C. pirolata was known to occur on spruce in Colorado, Montana, Oregon (Ar- thur 1934), and Washington (Shaw 1973). Apparent extension beyond the range of spruce may result from its perennial nature on the telial host and a possible lower fre- quency of critical requirements occurring for 'Intermountain Forest and Range Experiment Station, U.S. Department of Agriculture, Forest Service, Ogden, Utah, located at Shrub Sciences Labora- tory, Provo, Utah 84601. ^Rocky Mountain Forest and Range Experiment Station, U.S. Department of Agriculture, Forest Service, Forestry Sciences Laboratory, Arizona State University, Tempe, Arizona 8528L 262 Tune 1982 Nelson, Krebill: Cone Rust on Spruce 263 infection of spruce than for pyrola. Although unclear from the literature, however, it ap- pears that collection locations of the rust on Pyrola and Moneses spp. in the West are within the distributional range of spruce. Presently a clear extension beyond the range of spruce is the occurrence in Guatemala on Pyrola secunda L. (Cummins 1943), a great distance from the nearest indigenous spruce in northern Mexico (Martinez 1963). During the Pleistocene, spruce extended farther south and the fungus may have survived on pyrola to the present, when spruce died out, rather than spreading southward on pyrola. Inaccessibility of rusted cones to the casual collector is likely a factor in the apparent low frequency of this rust on spruce. Also, there are no specific, systematic annual sur- veys made in the western United States to de- tect rust diseases as there are in Canada. Little knowledge of the epidemiology of this fungus exists. Clinton and McCormick (1919), in Petri dish culture tests, failed to ob- tain infection of excised pyrola leaves using uredinial inoculum, even though success was achieved with other rust fungi. Eraser (1911, 1912, 1925) made a series of phenological ob- servations and inoculation studies of the rust in Nova Scotia and Saskatchewan, Canada. He observed that uredinia on pyrola matured and began releasing spores by early May, and telia germinated in late May. Pistillate cones were opening on spruce in the vicinity. In early July, the pycnial state was evident with a yellowing of cone scales and a yellow-col- ored resin flow. Several controlled field and laboratory inoculation tests indicated that about 48 hours of moist environment was suf- ficient for infection of cones, although the number of infected cones was low. Environmental requirements for infection of pyrola from urediniospores and aecio- spores are probably more frequently met than for spruce cones from basidiospores be- cause of the differing microenvironment of the host organs. Pyrola inhabits moist sites in the shade of dense tree stands, compared to the exposed tops of spruce trees. This differ- ence would be greater in semiarid regions of the West, and perhaps less so in the Pacific Northwest. Many species of Pyrola and two species of Moneses are known hosts of C. pirolata (Ar- thur 1934, Brown 1956, Shaw 1973). Damage to these species by the disease is apparently minor, with some atrophy and yellowing of leaves (Rice 1927, 1935). Disease symptoms reported on spruce cones in Canada include an early yellowing of cone scales, resin flow, premature browning, and— following aecial formation— a premature opening of the cones (Eraser 1912, Ziller 1974). Distortion or at- rophy of cones is not indicated in the liter- ature. Except for Picea breweriana S. Wats., Picea chihitahuana Martinez, and Picea rnexi cana Martinez, all native North American spruce species are known hosts of C. pirolata (Arthur and Kern 1906, Ziller 1957, Can. Dept. Environ. 1951-1975). Hedgcock (1912) described infected cones of Engelmann spruce {Picea engelmannii Parry) as aborted, with the only apparent damage being a re- duction in seed crop. The infection is thought to spread completely throughout the cone (Savile 1950). Damage to seeds in rusted cones apparently can be severe. Rhoads et al. (1918) indicate that no seeds are produced in infected cones. In the United States (Zasada and Gregory 1969) and in Canada (Can. Dept. Environ. 1951-1975), reports indicate that no sound seeds are produced, although no germination studies were reported to sup- port this. Eide (1927) in Norway found that 33 percent of the seeds from rusted cones germinated compared to 56.5 percent from nonrusted cones. Neger (1924) described dis- eased cones as forming little or no seed. In British Columbia, Sutherland (1981) found that the effect of cone rust on Picea glauca (Moench) Voss seed was to reduce yield, weight, and in some cases, the germinative capacity. Sporadic, relatively localized epidemics appear to characterize the occurrence of C. pirolata spruce cone rust in the contiguous western United States. In contrast, in Alaska, Zasada and Gregory (1969) reported the rust on white spruce over an extensive area south of the Alaska range in 1960, and again in 1968 north of the Alaska range near Fair- banks. Ziller (1974) attributes an "A" damage rating ("causes great or significant damage") to inland spruce cone rust (caused by C. piro- lata) in Canada. Damage ranging from light to nearly the entire cone crop destroyed in certain localities is reported regularly in the 264 Great Basin Naturalist Vol. 42, No. 2 various provinces of Canada (Can. Dept. En- viron. 1951-1975). Similar severity of dam- age occm-s in Norway (Jo Cones open Q Cones closed # Pyrola rust site 0000000#4 • ••••••OO ♦ IS o PF o UR NS ♦ ♦ ♦ o o UR o o o o ♦ o o ♦ ♦ ♦ o ♦ ♦ o o o 1 — ' — \ — ' — \ — ' — \ — ' — \ — ' — \ — ' — \ — ^ — I — ^ — h / 3 5 7 9 11 13 15 17 STUDY PLOT NUMBER (1950-2470 meters elev.) Fig. 3. Phenology of spruce ovulate cones and Chrysomyxa piwlata on Pijrola spp. in Huntington Canyon. June 1982 Nelson, Krebill: Cone Rust on Spruce 267 where the rust was found on pyrola. The an- nual checks from 1970 to 1978 of the area where the 1969 cone rust epidemic occurred failed to reveal the recurrence of any cone rust. Cones were present each year in varying amounts. Phenology of spruce cone rust.— By 25 May of the year studied, ovulate cones were beginning to open at the lowest elevations of Huntington Canyon, but the strobilate buds were still dormant at 2200 m and above, throughout the pyrola zone (Fig. 3). Incipient uredinial-telial sori were visible on the lower leaf surfaces of pyrola near Plot 10. By 14 June, cones had opened and then closed to the 2100 m level and were open from 2200 m to 2300 m in the lower pyrola zone. A few uredinial peridia were rupturing with some spore dispersal in progress. Telia were pres- ent at the base of sori, but there was no evi- dence of germination. By 29 June, cones had closed at only slightly higher elevation, and cones were open throughout the upper can- yon. Urediniospore dispersal was at a peak; teliospore formation was very sparse, and there was no evidence of germination. By 15 July, all spruce cones were long closed to the Huntington Canyon (Upper) o o Eph raim (Low e r } o o 50-- 20-- '^ 10 Spruce Cone Rust Epidemic — ^ 22-Year Mean highest plot. Pyrola leaves with rust sori had dried up, and there was no evidence of telial germination. Ovulate cones opened and closed within 20 days in the lower canyon. Sponilation of ure- dinial-telial sori lasted for 15-20 days begin- ning in mid-June. In 1969, aeciospore dis- persal lasted from late August to at least through September. Associated precipitation.— A com- parison of the yearly summer rainfall at Eph- raim and Huntington canyons (Fig. 4) re- vealed a close correlation in annual fluctuations (correlation coefficient [r = 0.82]). The mean annual summer rainfall at Ephraim was 12.4 cm compared to 24.8 cm at Huntington Canyon, with similar de- viations from the mean. During the 6 years preceding 1969 (the only year of known spruce cone rust outbreak), the annual pre- cipitation at the Huntington Canyon site was above or near the 22-year mean, with no ex- treme years on the low side (Fig. 4). Follow- ing 1969, there was a downtrend with some near-mean years, but most were below the mean. The frequency of extended spring and fall rainy periods and precipitation totals at "Regression Correlation r = 0.82 10 20 30 40 HUNTINGTON CANYON PRECIR cm A-/ V- -V -"^ v \ / V H \ ' 56 58 60 62 64 66 68 70 72 74 76 78 ANNUAL SUMMER PRECIPITATION (April -Sept J 1956-1977 Fig. 4. Comparison of summer rainfall at Ephraim and Huntington canyons from 1956 through 197 268 Great Basin Naturalist Vol. 42, No. 2 Ephraim follow a similar pattern (Fig. 5). A continuity of extended periods of rainy weather during spring and fall occurred at Ephraim during the 4 years preceding 1969 that did not occur in the following 8 years. Years with similar extended rainy periods fol- lowing 1969 (1970, 1972, 1973, 1975) were interrupted by dry seasons. Effect of spruce cone rust.— Our obser- vations of Chrysomyxa pirolata cone-rust symptoms were hmited to mid-September, when aecia were well open. Depressed re- sinous areas were common on infected cones, usually on one side. The resinous areas ap- peared to have developed more slowly or in- completely, resulting in a slight twisting of the cone. Cone scales opened prematurely on rusted compared to nonmsted cones, and were often twisted and malformed (Fig. 6). Not all infected cones appeared to be com- pletely infected; or at least in some parts of the cone, no aecial development occurred, and cone scales did not open prematurely. These areas were usually on the upper por- tion of cones (Fig. 7). Aecia formed primarily in a zone peripheral to the seeds (Fig. 8). An evaluation (Table 1) revealed that non- rusted cones yielded an average of 204 ap- parently sound seeds, and rusted cones 188. Seeds likely to disperse readily averaged 113 for nonmsted and 13 for rusted. A check of overwintered cones (12 nonmsted, 9 rusted) at the Huntington Canyon site the following summer showed an average of 18 seeds per cone remaining in nonmsted cones and 86 in rusted cones. Viability of seeds extracted from cones as determined by germination tests is indicated in Table 2. Of seeds dis- lodged by tapping cones, 71.0 percent germi- nated from nonrusted cones compared to 48.9 percent from msted cones. Remaining seed extracted by breaking cones apart ger- minated at 53.3 percent for nonmsted and 34.8 percent for rusted. Discussion and Conclusions Occurrence, habitat, and phenology.— In Huntington Canyon located in central Utah's high plateau country, Chrysomyxa pirolata cone rust of blue spruce was ob- served in a rare epidemic phase. Cone mst 5 7 I' S5 3 5 S2 tota I 65 ' 66 ' 67 ' 68 * 69 ^ 70 ' 7? ^ 72 ^ 73 ^ 74 ^ 75 ^ 76 ^ 77 YEARS of RECORD 1965-1977 Fig. ,5. Extended hourly rainy periods and spring and fall precipitation at Ephraim, Utah. June 1982 Nelson, Krebill: Cone Rust on Spruce 269 was limited to the spruce-fir zone where Py- rola spp. also occurred. Riparian blue spruce extended to lower elevations in the pinyon- juniper zone where neither cone infection nor pyrola were found. Cones were present on spruce above and below the cone rust zone. Studies by Eraser (1911, 1912, 1925) in- dicate that pistillate cones are susceptible to infection when cone scales open for pollina- tion; however, the precise period of suscep- tibility has not been established. In our study, the period of cone opening observed was closely correlated with the onset of uredinial- telial sporulation. During the single season observed, cones at lower elevations opened and closed 20 days or less prior to when the rust fungus began sporulation. Phenological progress in cone development with elevation, it appears, could limit the distance of in- fection from a point inoculum source. In Norway (Roll-Hansen 1967), the greatest in- cidence of cone rust was found at higher ele- vations and more northern latitudes; how- ever, abundant infection of pyrola occurred elsewhere. The cause was attributed to unfa- vorable climatic conditions for infection of spruce cones in these areas. Chrysomyxa piro- lata cone rust is rare in Canadian Pacific Coast areas compared to other parts of west- em Canada (Ziller 1974). Climate and telial hosts did not appear to be limiting since the closely related Chrysornyxa monesis Ziller oc- curs on Sitka spruce {Picea sitchensis [Bong.] Carr.) cones and Moneses uniflora (L.) A. Gray in coastal areas (Ziller 1974). Our study did not include investigation of variation in duration of favorable moisture during precip- itation periods with elevation. However, evi- dence suggests that host phenology as well as habitat as influenced by elevation is a factor in the progressively narrower niches of the spruce cone-rust fimgus proceeding south- ward across the continent from northern lati- tudes. The uredinial-telial sporulation period lasted for 15-20 days, beginning in mid- June. The 1972 spring season was later than nor- mal, and therefore spore dispersal could oc- cur from May through June into early July. Teliospores did not develop fully, and tiiere was no germination evident. This result was likely because of the dry spring of 1972 (Figs. 4 and 5). This also may have speeded the death of pyrola leaves upon which sori formed. Fraser (1911), in eastern Canada, and Rice (1927), in the New England states, ob- served that teliospore formation followed urediniospore formation by several weeks. Although there was poor telial formation in our study, it appeared to be almost simulta- neous with uredinial formation as Savile (1950) reported, and is typical of the Chryso- myxa rust fungi in general (Ziller 1974). Peri- ods of rainy weather may be necessary for abundant teliospore formation. Even though the aecial-pycnial stage on spruce is apparently relatively rare in the southern distribution of this cone rust on the North American Continent, it is significant that the sexual stage does occur. There is then the chance for genetic recombination and diversification of the rust fungus without dependence on migration of genes from northern latitudes on Pyrola. Associated precipitation.— The close correlation of Huntington Canyon and Eph- raim summer precipitation fluctuations al- lowed some confidence in projecting the Ephraim hourly record. Other than elevation, most differences are probably accounted for by short-term thunderstorms. Duration of ex- tended rain at Huntington Canyon, however, was likely longer because the total precipi- tation was nearly twice the amount at Eph- raim. Rainy periods of four hours or more in spring and fall were infrequent in the 13- year-record studied (Fig. 5), and therefore the Table 2. Effect of Chrysomyxa pirolata cone rust on Table 1. Effect of Chrysornyxa pirolata cone rust on viability of blue spruce seed. blue spruce s< ;ea yieic 3 ana aispersa . No. seeds' % Germination No. cones' Average no. seeds per cone Nonrusted Rusted Nonrusted Rusted Dislodged by tapping Removed by breaking Total Dislodged by tapping Removed by breaking 600 600 325 600 71.0 48.9 53.3 34.8 Nonrusted Rusted 89 48 113 13 91 204 175 188 'Total cones from six plots. ■Maximum 100 seeds per plot. 270 Great Basin Naturalist Vol. 42, No. 2 coincidence of susceptible cones, favorable moisture, and fungus inoculum is also prob- ably infrequent. Based on spring rainy peri- ods, the chance for favorable moisture for in- fection during springs of 1970, 1973, and 1975 appears just as positive as 1969, al- though no cone infection was detected. Perhaps, then, a continuity of extended spring and fall rains over several years is nec- essary for an epidemic. With dry seasonal in- terruptions, the fungus population could de- cline to levels that do not permit detectable amounts of cone infection even though spring rains during some years may be favorable. There is, however, no cause-and-effect rela- tionship established here. There is not enough reliable knowledge about the epide- miology of C. pirolata cone rust to relate epi- demics to weather records in a precise sense. The urediniospores produced on the Pyrola spp. observed in this study adhered rather strikingly, suggesting that dispersal may be primarily by rain splash, thus restricting in- tensification of the rust in the uredinial phase to the immediate locality. Widespread dis- persal of the fungus to pyrola would then seem to be more dependent on aeciospores that are readily dislodged from cones over a long period. If these possibilities are true, it would lend support to the importance of fall rainy periods. Because of the systemic-per- ennial nature of the pyrola rust infection, an important factor in the need for consecutive years of favorable moisture would be the number of years rusted pyrola plants live. Effect of spruce cone rust.— Symptoms of C. pirolata cone rust of Colorado blue spruce observed in this study wer^ somewhat different from what is reported in the liter- ature for other species of spruce. Depressed resinous areas on infected cones appeared to cause slight twisting of the cones. Portions of infected cones appeared to be rust free, or at least there were areas where no aecia formed. Aecial pustules formed primarily in a zone peripheral to the seeds, which may be related to the amount of seed damage. The disease appeared to have only a slight effect on development of seeds. There was about a 25 percent reduction in seed germinability with seed from rusted cones. This indicates that cone rust of blue spruce is less damaging to seed development than reported for other species of spruce. Relatively few seeds could Fig. 6. Chrysomyxa pirohta rust-diseased Colorado blue spruce cones (three left) and nonrusted cones (three right). Note the twisted, malformed nature of the rusted compared to nonrusted cones. June 1982 1 Nelson, Kkebill: Cone Rust on Spruce 271 ^^^HL>«^^^^ TK i^^ K' : ^9|^ j ■b' Fig. 7. Rusted (left) and nonnisted (right) cones of blue spruce. Note tfiat cone scales on upper half of rusted cone have not opened. be easily dislodged artificially from rusted cones compared to nonrusted. This indicates that the disease severely impedes seed dis- persal; and this was substantiated upon exam- ination of naturally overwintered cones. Ae- cial spore masses, resinosus, and malformation of cone scales prevented seed dispersal to the extent that the seed crop was effectually destroyed. Literature Cited Arthur, J. C. 1907-1931. Order Uredinales. Pages 1-969 1)1 North America Flora. Vol. 7, Part 2-1.3. New York Botanical Gardens, New York. 1934. Manual of the rusts in the United States and Canada. Purdue Research Foundation, La- fayette, Indiana. 438 pp. Arthur, J. C, and F. D. Kern. 1906. North American species of Peridermiiim. Bull. Torrey Bot. Club 33:403-438. Fig. 8. Rust-infected blue spruce cones in cross sec- tion. Note that "white" spore masses are peripheral to the inner seed zone. Brown, A. M. 1956. A checklist of plant rusts in Can- ada. Canada Dept. of Agric. Publ. 976. 51 pp. Canada Department of Environment. 1951-1975. An- nual Report, Forest Lisect and Disease Survey, Canadian For. Serv., Queen's Printer, Ottawa. Cash, E. K. 1953. A checklist of Alaskan fimgi. Plant Dis. Reptr. Suppl. 219. 70 pp. Clinton, G. R., and F. A. McCormick. 1919. Rust in- fection on leaves in Petri dishes. Conn. Agric. Expt. Sta. Bull. 260:475-501. Cummins, G. B. 1943. Annotated checklist and host in- dex of the rusts of Guatemala. Plant Dis. Reptr. Suppl. 142. 131 pp. EiDE, E. 1927. Underscikelse av nordenfjelsk granfrcS 1925. Med. fra Det Norske Skogforsciksvesen, H. 8:15-39. Eraser, W. P. 1911. Cultures of some heteroecious rusts. Mycologia 3:67-74. 1912. Cultures of heteroecious rusts. Mycologia 4:175-193. 1925. Culture experiments with heteroecious rusts in 1922, 1923, and 1924. Mycologia 17:78-86. 272 Great Basin Naturalist Vol. 42, No. 2 Gaumann, E. 1959. Die Rostpilze Mitteleuropas, Bul- cher Co., Bern. Beitrage zur Kryptogamenflora der Schweiz, Bd. 12. GiLBERTSON, R. L., AND J. McHenry. 1969. Checklist and host index for Arizona rust fungi. Univ. of Ariz. Agric. Expt. Sta. Tech. Bull. 186. 40 pp. Hedgcock, G. G. 1912. Notes on some western Ure- dineae which attack forest trees. Mycologia 4:141-147. JdRSTAD, I. 1935. Uredinales and Ustilaginales of Trcindelag. Det Kyi Norske Videnskabers Selskabs Skrifler NR 38. 90 pp. 1940. Uredinales of northern Norway. Skr. utg. av. Det. Norske Vindensk.-Akad. i, Oslo. I. Mat.- Naturv. Klasse. 1940. No. 6. 145 pp. KiMMEY, J. W., AND J. A. Stevenson. 1957. A forest dis- ease survey of Alaska. Plant Dis. Reptr. Suppl. 247. pp. 87-98. Martinez, M. 1963. Las Pinaceas Mexicanas, 3d ed. Universidad Nacional Autonoma de Mexico, Mexico City. 401 pp. Neger, F. W. 1924. Die krankheiten unserer Waldbaume und der wichtigsten Gartengeholze. 2d ed. Ferdinand Enke, Stuttgart. 296 pp. Nelson, D. L., and R. G. Krebill. 1970. Effect of Chrysomyxa pirolata cone rust on dispersal and viability of Picea pungens seeds. Phytopathology Abstr. 60:1305. Peterson, R. S. 1961. Notes on western rust fungi. I. Chrysomyxa. Mycologia 53:427-431. Rhoads, a. S., G. G. Hedgcock, E. Bethel, and C. Hartley. 1918. Host relationships of the North American rusts other than Gymnosporangiums, which attack conifers. Phytopathology 8:309-352. Rice, M. A. 1927. The haustoria of certain rusts and the relation between host and pathogen. Bull. Torrey Bot. Club 54:63-153. 19.35. The cytology of host-parasite relations. Bot. Rev. 1:327-354. Roll-Hansen, F. 1967. On diseases and pathogens of forest trees in Norway, 1960-1965. (Om sykdom- mer og patogener pa skograernei Norge, 1960-1965). Saertrykk av Meddelelser fra Det Norske Skogforseiksvesen. Vollebekk, Norge. Nr. 80, Bind XXI, pp. 174-246. Savile, D. B. O. 1950. North American species of Chrysomyxa. Canadian Jour. Resc. 28:318-330. 1953. Short-season adaptations in the rust fimgi. Mycologia 45:75-87. Shaw, C. G. 1973. Host fungus index for the Pacific Northwest— I. Hosts;— II. Fungi. Washington Agr- ic. Expt. Sta. Bull. 765, 766. Sutherland, J. R. 1981. Effects of inland spruce cone rust, Chrysomyxa pirolata Wint., on seed yield, weight and germination. Canadian For. Serv., Res. Notes l(2):8-9. Zasada, J. C, and R. a. Gregory. 1969. Regeneration of white spruce with reference to interior Alaska: a literature review. USDA For. Serv. Res. Pap. PNW-79, 37 pp. ZiLLER, W. G. 1957. Fungi of British Cokmibia depos- ited in the herbarium of the Forest Biology Labo- ratory, Victoria, BC, Canada. For. Biol. Div., Sci. Serv., Agric. Dept., Ottawa, Canada. 1974. The tree rusts of western Canada. Canada Dept. 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No. 6 The bark and ambrosia beetles of North and Central America (Coleoptera: Scolytidae), a taxonomic monograph. $60. TABLE OF CONTENTS Utah plant types-historical perspective 1840 to 1981-annotated list, and bibliogra- phy. Stanley L. Welsh 229 A new species of Cryptantha (Boraginaceae) from Nevada. Kaye H. Thorne and Lar- ry C. Higgins ^gg New taxa of thistles {Cirsium: Asteraceae) in Utah. Stanley L. Welsh 199 A species of Cryptantha (Boraginaceae) dedicated to the memory of F. Creutzfeldt. Stanley L. Welsh 203 Algal populations in Bottle Hollow Reservoir, Duchesne County, Utah. Jeffrey Jo- hansen, Samuel R. Rushforth, and Irena Kaczmarska 205 Herpetological notes from the Nevada Test Site. Wilmer W. Tanner 219 New species of American bark beetles (Coleoptera: Scolytidae). Stephen L. Wood 223 Temperature-related behavior of some migrant birds in the desert. George T. Austin and J. Scott Miller 232 Description of a new Phalacwpsylla and notes on P. alios (Siphonaptera: Hys- trichopsyllidae). R. B. Eads and E. G. Campos 241 Vegetation of the mima mounds of Kalsow Prairie, Iowa. Jack D. Brotherson 246 Occurrence and effect of Chrysomyxa piwlata cone rest on Picea pungens in Utah. David L. Nelson and Richard G. Krebill 262 THE GREAT BASIN NATURALIST Volume 42 No. 3 September 30, 1982 Brigham Young University MUS. CO MP. ZOOL LIBRARY FEB - 2 H UNIVERSITY 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). 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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. 11-82 650 61360 ISSN 017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 42 September 30, 1982 No. 3 BUCCAL FLOOR OF REPTILES, A SUMMARY Wilmer W. Tanner' and David F. Avery^ Abstract.— A general survey of the information presently available on the osteology and myology of the hyobran- chial apparatus. Included in the survey are examples of the hyobranchial skeleton of the major groups of reptiles, in- cluding the Chelonia, Crocodilia, Rhynchocephalia, and Squamata. The myology treats the muscles directly associ- ated with the hyoid as well as those associated with the functioning of the apparatus, but not arising or inserted directly on or from the hyoid. The innervation of the hyobranchial apparatus is reviewed and briefly discussed based on the information available in a few major studies. An attempt is made to cite all pertinent literature references, and in Tables 1 and 2 the references to basic areas are indicated. Twenty-nine plates and figures are included, some of which represent original research. I. Introduction Few anatomical areas have been subjected to such pronounced evolutionary changes as have the branchial apparatus and its deriva- tives in the vertebrate series. The hyoid ap- paratus has responded to these numerous adaptive changes with structural and func- tional modifications. One needs only to con- template the change necessary in adapting from a structure bearing gills to one associ- ated with lungs, from an immovable to a highly flexible tongue, or to the development of a lamyx and archaic voice to appreciate the anatomical importance of this area. Fur- thermore, the class Reptilia consists of both primitive (turtles, crocodilians, and Spheno- don) and specialized (lizards and snakes) forms that include organisms possessing con- siderable structural diversification. In reptiles the buccal floor consists of os- seous and cartilaginous elements of the bran- chial skeleton and the associated connective and muscular tissues. Included among the skeletal elements are the jaws, hyoid appa- ratus, laryngeal cartilages, and tracheal rings. The associated fleshy parts include the hypo- branchial throat musculature, the tongue, and the nerves and blood vessels associated with them. There is also a variety of glands associ- ated with the buccal floor; these are usually -involved with the production of saliva that may be poisonous. A complete comparative anatomical treat- ise on the buccal floor is not possible at this time, primarily because the necessary infor- mation is not available. Some anatomical studies on reptiles are precise and show con- siderable detail; however, the studies have too often been concerned primarily with one series of bones or one group of muscles rather than an entire anatomical pattern. As a re- sult, we will confine our remarks to the pres- ent knowledge of the hyoid structure and as- sociated muscles and nerves in the floor of the reptilian mouth. Many studies touch on the subject at hand in various ways. We have, therefore, included in the bibliography many 'Life Science Museum, Brigham Young University, Provo, Utah 84602. 'Department of Biology, Southern Connecticut State College, New Haven, Cormecticut 06515. 273 274 Great Basin Naturalist Vol. 42, No. 3 studies not cited in the text. These have been useful in our examination of the materials available and are as follows: Adams 1919, 1925, Ashley 1955, Barrows and Smith 1947, Beddard 1905, Bellairs 1950, Bergman 1961, 1965, Boltt and Ewer 1964, Brock 1938, Bull- ock and Tanner 1966, Byerly 1926, Chaine 1902, Chiasson 1962, Cowan and Hick 1951, Davis 1934, Duda 1965, Dullemeijer 1956, 1958, El Toubi 1938, 1947a, 1947b, El Toubi and Kalil 1952, Eyal-Giladi 1964, Evans 1955, Gandolfi 1908, Gans 1961, George 1948, George and Shad 1954, 1955, Haas 1952, 1960, 1968, 1973, Harris 1963, Hey- mans 1970, lordansky 1970, Iyer 1942, 1943, Kamal, Hammouda, and Mokhtar 1970, Kes- teven 1944, Kingman 1932, Kluge 1962, Kochva 1958, Liem, Marx, and Rabb 1971, Mahendra 1949, Malam 1941, McKay 1889, Minot 1880, Mivart 1867, Norris and Lowe 1951, Oldham, Smith, and Miller 1970, Park- er 1880, Ping 1932, Presch 1971, Rathor 1969, Reese 1923, Rice 1920, Rieppel 1981, Rosenberg 1968, Sanders 1870, 1872, 1874, Schumacher 1956c, Sewertzoff 1929, Shah 1963, Sidky 1967, Siebenrock 1892a, 1892b, 1893, 1894, 1895, Sinitsin 1928, and Varkey 1979. Tables 1 and 2 provide additional informa- tion on the material covered by these and other authors dealing with buccal floor and associated structures. II. Hyoid Apparatus General The branchial skeleton, including the vis- ceral arches, which we have associated with the more primitive gill-bearing vertebrates, has been recast in the tetrapods where its structure and function have been modified. The branchial skeleton now appears in tetra- pods as a part of the skull; it includes the jaw and the hearing apparatus, as well as the la- rynx and trachial cartilage supports. The tetrapod has also retained the more central part of the old visceral skeleton, which is now known as the hyoid apparatus. Because reptiles have lost the gill appa- ratus in all stages of development, the hyoid apparatus has assumed the function of a sup- port for the tongue, glottis, and sometimes an extended dewlap. In modern reptiles, the hyoid is composed of several osseous and car- tilagenous elements and exhibits a variety of degrees of ossification. As a general rule, the larger (or older) the animal, the more ossified is the hyoid apparatus. In most reptiles, ex- cept in some snakes, the hyoid apparatus is a spreading, flexible structure that occupies space in, and forms a support for, most of the floor of the oropharynx. Although the phylogenetic relationships of the hyoid apparatus and visceral arches are not completely understood, it is known that the hyoid apparatus is derived from the hyoid cartilage and the two succeeding arches. Ro- mer (1956) believes that the hyoid of ances- tral reptiles must have been more extensive and that traces of a third branchial cornu can be seen in some reptilian embryos. The third cornu is well demonstrated in monotreme mammals. The nomenclature pertaining to the hyoid is not uniform. Furbringer (1922) describes the first two pairs of arches as the cornu hyale and the cornu brachiale I, respectively; the third arch is called the cornu branchiale II. This latter arch is referred to by Beddard (1907) as the branchial process and as the basibranchial by Gnanamuthu (1937). The third arch is seemingly absent in several rep- tiles, causing some workers to refer to the re- maining two arches as the anterior and poste- rior cornua. Unfortunately, the identity of the third arch has not been clearly ascer- tained. The third arch may be a degenerate structure expressed as projections from the basihyoid or body of the hyoid, or it may be present as a separate arch with either the first or second arch being lost. In the Ophidia and some burrowing lizards such as Anniella, Dibamus, Acontias, Acontophiops, and Typh- losaurus, the hyoid is greatly reduced and the identity of the posterior cornua is not posi- tively established. (See Rieppel 1981 for a more complete discussion.) A similar situa- tion exists in the Testudines and Crocodilia. The development of the hyoid apparatus has been discussed by Rathke (1839), Kallius (1901), Howes and Swinnerton (1901), Peyer (1912), Edgeworth (1935), DeBeer (1937), Pringle (1954), El Toubi and Kamal (1959a,b), El Toubi and Majid (1961), Kamal and Hammouda (1965), Langebartel (1968), Rieppel (1981), and others (Table 1). These September 1982 Tanner, Avery: Buccal Floor of Reptiles 275 Table 1. Publications dealing with the buccal floor of reptiles. Genus Hyoid Tongue Order Chelonia Suborder Pleurodina Musculature Nerves Pelomedusidae Pehtsios Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Chelidae Batrochemys CJielodina Furbringer 1922 Winokur 1974 Poglayen-Neuwall 1953 Graper 1932 Kesteren 1944 Poglayen-Neuwall 1953 Shah 1963 Poglayen-Neuwall 1953 Kesteren 1944 Poglayen-Neuwall 1953 Suborder Cryptodira Dermatemydidae Dennatemys Furbringer 1922 Chelydridae Chelydra Kinosternon Sternotherus Furbringer 1922 Edgeworth 1935 Schumacher 1973 Furbringer 1922 Schumacher 1973 Furbringer 1922 Schumacher 1973 Winokur 1974 Camp 1923 Graper 1932 Poglayen-N eu wall 1953 Schumacher 1973 Poglayen-Neuwall 1953 Schumacher 1973 Poglayen-Neuwall 1953 Schumacher 1973 Poglayen-Neuwall 1953 Soliman 1964 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Testudinidae Chrysemys C/em 771 1/5 Cuora Deirochelys Dermaiemys Emys Furbringer 1922 Ashley 1955 Schumacher 1973 Siebenrock 1898 Furbringer 1922 Schumacher 1973 Furbringer 1922 Furbringer 1922 Walter 1887 Furbringer 1922 Schumacher 1973 Winokur- Pers. Comm. Sewentzoff 1929 Poglayen-Neuwall 1953 Ashley 1955 Schumacher 1973 Graper 1932 Lubosch 1933 Schumacher 1973 Poglayen-Neuwall 1953 Shah 1963 Walter 1887 Schumacher 1973 Poglayen-Neuwall 1953 Lubosch 1933 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Gopherus Winokur 1973 George & Shad 1955 276 Great Basin Naturalist Vol. 42, No. 3 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Graptemys Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Geochelone (Testudo) Bojanus 1819 Furbringer 1922 Edgeworth 1935 Hacker & Schumacher 1955 Schumacher 1973 Bojanus 1819 Bojanus 1819 Graper 1932 Edgeworth 1935 Lubosch 1933 Poglayen-Neuwall 1953 Schumacher 1973 Lubosch 1933 Poglayen-Neuwall 1953 Malachemys Psetidemys Furbringer 1922 Schumacher 1973 Poglayen-Neuwall 1953 Ashley 1955 Poglayen-Neuwall 1953 Schumacher 1973 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Terrapene Trionychidae Furbringer 1922 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Trionyx {Amy da) Siebenrock 1898 Sondhi 1958 Furbringer 1922 Schumacher 1973 Sondhi 1958 Graper 1932 Lubosch 1933 Poglayen-Neuwall 1953 Schumacher 1973 Poglayen-Neuwall 1953 Lissemys Cheloniidae Caretta Demiachelyidae Dermochelys Furbringer 1922 Sondhi 1958 Schumacher 1973 Furbringer 1922 Schumacher 1973 Schumacher 1973 Gnananuthu 1937 Sondhi 1958 Order Rhynchocephalia Sphenodontidae Sphenodon Osawa 1898 Sewertzoff 1929 Howes & Swinnerton 1901 Furbringer 1922 Edgeworth 1931,35 Rieppel 1978 George & Shad 1954 Sondhi 1958 Schumacher 1973 Poglayen-Neuwall 1953 Schumacher 1973 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953/54 Schumacher 1973 Osawa 1898 Camp 1923 Byerly 1926 Edgeworth 1931,35 Lightoller 1939 Kesteven 1944 Rieppel 1978 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953 Poglayen-Neuwall 1953/54 Osawa 1898 Lubosch 1933 Kesteven 1944 Rieppel 1978 September 1982 Tanner, Avery: Buccal Floor of Reptiles 277 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Order Squamata Suborder Sauna Gekkonidae Ascolabotes Cneniospis Coleonyx Eiiblepharis Gehrydra Gekko Gijmnodactijlus Hemidactijlus Phyllodactyhts Platydactyhis Ptychozoon Stenodactylus Tarentoh Thecodactylus Uroplatus Dibamidae Dibamiis Iguanidae Ambryrhynchus Anolis Basiliscus Brachylophiis Callisaurus Chalarodon Richter 1933 Camp 1923 Kluge 1962 Cope 1892 Camp 1923 Richter 1933 Camp 1923 Richter 1933 Richter- 1933 Zavattarl 1908 Richter 1933 Edgeworth 1935 Cope 1892 Richter 1933 Richter 1933 Verslvys 1898, 1904 Camp'l923 Edgeworth 1935 Rieppel 1981 Avery & Tanner 1971 Cope 1892 Zavattari 1908 Camp 1923 Avery & Tanner 1971 Cox & Tanner 1977 Avery & Tanner 1971 Sewertzoff 1929 Ping 1932 Avery & Tanner 1971 Gnanamuthu 1937 Avery & Tanner 1971 Avery & Tanner 1971 Camp 1923 Edgeworth 1935 Camp 1923 Camp 1923 Lubosch 1933 Brock 1938 Kesteven 1944 Zavattari 1909 Ping 1932 Edgeworth 1935 Gnanamuthu 1937 Sanders 1870 Poglayen-Neuwall 1954 Gnanamuthu 1937 Poglayen-Neuwall 1954 Kesteven 1944 Case 1968 Avery & Tanner 1971 Kesteven 1944 Gnanamuthu 1937 Camp 1923 Avery & Tanner 1971 Cox & Tanner 1977 Avery & Tanner 1971 Lubosch 1933 Kesteven 1944 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 Kesteven 1944 Willard 1918 Kesteven 1944 Renous-Lecuru 1972 278 Great Basin Naturalist Vol. 42, No. 3 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Chamaeleolis Beddard 1907 Conolophus Avery & Tanner 1971 Avery & Tanner 1971 Avery & Tanner 1971 Coplwsaunis Cox & Tanner 1977 Cox & Tanner 1977 Crotaphijtus Cope 1892 Robison & Tanner 1962 Davis 1934 Robison & Tanner 1962 Ctenosaura Oelrich 1951 Avery & Tanner 1971 Oelrich 1956 Avery & Tanner 1971 Oelrich 1956 Avery & Tanner 1971 Cychira Avery & Tanner 1971 Gandolfi 1908 Avery & Tanner 1971 Avery & Tanner 1971 Dipsosaurus Cope 1892 Avery & Tanner 1971 Avery & Tanner 1971 Avery & Tanner 1971 Enifaliosaurus Avery & Tanner 1971 Avery & Tanner 1971 Avery & Tanner 1971 Holbrookia Cox & Tanner 1977 Cox & Tanner 1977 Iguana Edgeworth 1935 Avery & Tanner 1971 Oldham & Smith 1975 Gandolfi 1908 Avery & Tanner 1971 Oldham & Smith 1975 Mivart 1867 Edgeworth 1935 Poglayen-Neuwall 1954 Avery & Tanner 1971 Oldham & Smith 1975 Poglayen-Neuwall 1954 Oldham & Smith 1975 Liolaemus Gandolfi 1908 Opiums Avery 6c Tanner 1971 Avery & Tanner 1971 Avery & Tanner 1971 Phrynosoma Cope 1892 Camp 1923 Richter 1933 Jenkins & Tanner 1968 Sanders 1874 Camp 1923 Jenkins & Tanner 1968 Pohjchrus Richter 1933 Sauroinalus Avery & Tanner 1964, 1971 Avery & Tanner 1971 Avery & Tanner 1964, 1971 Sceloporus Cope 1892 Secoy 1971 Tropidurus Zavattari 1908 Edgeworth 1935 Zavattari 1908 Edgeworth 1935 Uma Cox & Tanner 1977 Cox & Tanner 1977 Urosaurus Fanghella, Avery & Tanner 1975 Fanghella, Avery & Tanner 1975 Uta Fanghella, Avery & Tanner 1975 Fanghella, Avery & Tanner 1975 September 1982 Tanner, Avery: Buccal Floor of Reptiles 279 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Agamidae Agama Amphibolurus Calotes Ceratophora Ch lam ydosa u rus Cophotis Draco Hydrosaurus Leiolepis Lyriocephalus Otocryptis Phrynocephalus Physignathus Sitana Uromastix Edgeworth 1935 El-Toubi 1947 Harris 1963 Eyal-Giladi 1964 Richter 1933 Zavattari 1908 Camp 1923 Richter 1933 Edgeworth 1935 Iyer 1943 Richter 1933 Beddard 1905 Richter 1933 Richter 1933 Richter 1933 Richter 1933 Richter 1933 Richter 1933 Richter 1933 Kesteven 1944 Kesteven 1944 Islam 1955 Tilak 1964a,b Gandolfi 1908 Gandolfi 1908 Gandolfi 1908 Sewerteoff 1929 Gnanamuthu 1937 Gnanamuthu 1937 Sewertzoff 1929 DeVis 1883 Lubosch 1933 Edgeworth 1935 Poglayen-Neuwall 1954 Harris 1963 Poglayen-Neuwall 1954 Camp 1923 Gnanamuthu 1937 Poglayen-Neuwall 1954 DeVis 1883 Gnanamuthu 1937 Sanders 1872 Poglayen-Neuwall 1954 Kesteven 1944 Kesteven 1944 Gnanamuthu 1937 Furbringer 1922 Lubosch 1933 Edgeworth 1935 George 1948 Poglayen-Neuwall 1954 Throckmorton 1978 Lubosch 1933 Poglayen-Neuwall 1954 Carpenter et al. 1977 Poglayen-Neuwall 1954 Gnanamuthu 1937 Poglayen-Neuwall 1954 Renous & Lecuru 1972 Poglayen-Neuwall 1954 Kesteven 1944 Poglayen-Neuwall 1954 Chamaeleonidae Chamaeleo Zavattari 1908 Edgeworth 1935 Gnanamuthu 1937 Jollie 1960 Lubosch 1932 Gnanamuthu 1937 Mivart 1870 Mivart 1876 Zavattari 1908 Camp 1923 Lubosch 1933 Edgeworth 1935 Gnanamuthu 1937 Kesteven 1944 Poglayen-Neuwall 1954 Gnanamuthu 1937 Kesteven 1944 Poglayen-Neuwall 1954 Scincidae Ablepharus Sewertzoff 1929 280 Great Basin Naturalist Vol. 42, No. 3 Table 1 continued. Genus Hyoid Chalcides Richter 1933 El Toubi 1938 El Toubi & Kamal 1959a,b Eumeces Cope 1892 Zavattari 1908 Richter 1933 Nash & Tanner 1970 Lygosoma Richter 1933 Mabuya Richter 1933 Gnanamuthu 1937 Tao & Ramaswami 1952 Nessia Richter 1933 Riopa Richter 1933 Scincus Richter 1933 Tiligua Beddard 1907 Tongue Musculature Zavattari 1908 Edgeworth 1935 Nash & Tanner 1970 Gnanamuthu 1937 Gnanamuthu 1937 Lightoller 1939 Kesteven 1944 Poglayen-Neuwall 1954 Nerves Soliman & Hegazy 1971 Lightoller 1939 Poglayen-Neuwall 1954 Trachysaurus Beddard 1907 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 Typhlosaurus Rieppel 1981 Cordylidae Cordylus Beddard 1907 Camp 1923 Richter 1933 Edgeworth 1935 Camp 1923 Edgeworth 1935 Gerrhosaurus Camp 1923 Camp 1923 Zonrus Camp 1923 Lacertidae Acanthodactylus Richter 1933 Cabrita Gnanamuthu 1937 Gnanamuthu 1937 Lacerta Walter 1887 Zavattari 1908 Richter 1933 Edgeworth 1935 Sewertzoff 1929 Walter 1887 Camp 1923 Edgeworth 1935 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 Ophisops Richter 1933 Teiidae Ameiva Richter 1933 Fisher & Tanner 1970 Minot 1880 Sewertzoff 1929 Presch 1971 Poglayen-Neuwall 1954 Fisher & Tanner 1970 Poglayen-Neuwall 1954 September 1982 Tanner, Avery: Buccal Floor of Reptiles 281 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Cnemidophorus Neusticiirus Tupinainbis Anguinidae Angitis Gerrhonotus Ophiosaurus Xenosauridae Shinosaitms Cope 1892 Fisher & Tanner 1970 Richter 1933 Zavattari 1908 Reese 1932 Edgeworth 1935 Jollie 1960 Richter 1933 Walter 1887 Cope 1892 Walter 1887 Presch 1971 Sewertzoff 1929 Sewertzoff 1929 Poglayen-Neuwall 1954 Fisher & Tanner 1970 Zavattari 1908 Camp 1923 Edgeworth 1935 Poglayen-Neuwall 1954 Camp 1923 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 Poglayen-Neuwall 1954 McDowell & Bogart McDowell & Bogart Haas 1960 1954 1954 Xenosaiirus Helodermatidae Helodenna Varanidae Varaniis Lanthanotidae Lanthanotus Anniellidae Anniella Camp 1923 McDowell & Bogart Camp 1923 McDowell & Bogart 1954 Haas 1960 1954 Cope 1892 McDowell & Bogart 1954 Richter 1933 McDowell & Bogart 1954 Sondhi 1958 McDowell & Bogart 1954 McDowell 1972 Rieppel 1981 Cope 1892 Rieppel 1981 McDowell & Bogart 1954 Sewertzoff 1929 McDowell & Bogart 1954 Sondhi 1958 McDowell & Bogart 1954 Camp 1923 Poglayen-Neuwall 1954 Bradley 1903 Camp 1923 Edgeworth 1935 Gnanamuthu 1937 Lightoller 1939 Kesteven 1944 Poglayen-Neuwall 1954 Sondhi 1958 Poglayen-Neuwall 1954 Watkinson 1906 Lightoller 1939 Kesteven 1944 Poglayen-Neuwall 1954 Camp 1923 282 1 Great Basin Naturalist Vol. 42, No. 3 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Amphisbaenidae Amphisbaena Camp 1923 Richter 1933 Jollie 1960 Smalian 1885 Camp 1923 Anopsibaena Smalian 1885 Bipes Renous 1977 Smalian 1885 Blanus Smalian 1885 Monopeltis Richter 1933 Rhineura Cope 1892 Camp 1923 Trogonophis Smalian 1885 Xantusidae Xantusia Savage 1963 Cope 1892 Camp 1923 Suborder Serpentes Anomalepididae Anomalepis Smith & Warner 1948 Haas 1968 Helminthophis List 1966 Langebartel 1968 Langebartel 1968 Langebartel 1968 Liotyphlops List 1966 Langebartel 1968 Langebartel 1968 Langebartel 1968 Typhlopidae Typhhps List 1966 Langebartel 1968 Langebartel 1968 Langebartel 1968 Typhlophis Evans 1955 List 1966 Evans 1955 Evans 1955 Leptotyphlopidae Leptotyphhps Smith & Warner 1948 List 1966 Langebartel 1968 Oldham, Smith & Miller 1970 Langebartel 1968 Oldham, Smith & Miller 1970 Langebartel 1968 Uropeltidae Platyplectrurus Langebartel 1968 Langebartel 1968 Langebartel 1968 Plectrurus Rieppel 1981 Rhinophis Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Langebartel 1968 Silybura Langebartel 1968 Uropeltis Langebartel 1968 Langebartel 1968 September 1982 Tanner, Avery: Buccal Floor of Reptiles 283 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Aniliidae Anilius Smith & Warner 1948 Langebartel 1968 Rieppel 1981 Langebartel 1968 Langebartel 1968 Cylindrophis Smith &c Warner 1948 Langebartel 1968 Lubosch 1933 Langebartel 1968 Lubosch 1933 Langebartel 1968 Xenopeltidae Xenopeltis Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Langebartel 1968 Boidae Aspidites Smith & Warner 1948 Langebartel 1968 Boa Langebartel 1968 Gibson 1966 Calaharia Langebartel 1968 Langebartel 1968 Langebartel 1968 Charina Langebartel 1968 Langebartel 1968 Langebartel 1968 Chondropython Langebartel 1968 Constrictor Langebartel 1968 Langebartel 1968 Langebartel 1968 Entjgrus Langebartel 1968 Epicrates Langebartel 1968 Langebartel 1968 Langebartel 1968 Eryx Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Langebartel 1968 Eunectes Langebartel 1968 Anthony & Serra 1950 Langebartel 1968 Langebartel 1968 Liasis Langebartel 1968 Langebartel 1968 Langebartel 1968 Lichanura Langebartel 1968 Hershkowitz 1941 Loxocemus Smith & Warner 1948 Langebartel 1968 Nardoana Langebartel 1968 Python Furbringer 1922 Langebartel 1968 Oldham, Smith & Miller 1970 Lubosch 1933 Edgeworth 1935 Kesteven 1944 Frazzetta 1966 Lubosch 1933 Kesteven 1944 Langebartel 1968 Sanzinia Trachyboa Langebartel 1968 Langebartel 1968 Langebartel 1968 Oldham, Smith & Miller 1970 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 284 Great Basin Naturalist Vol. 42, No. 3 Table I continued. Genus Hyoid Tongue Musculature Nerves Colubridae Achalinus Achrochordiis Adelphicus Amblycephahis Aparallactus Apostolepis Atrethim Boiga Carphophis Cerberus Chersodroinus Chersydrus Chri/sopelea Clelia Coluber Coniophanes Conophis Conepsis Crotaphopehis Cyclagras Dasypeltis Dendrophidion Diadophis Dipsadotoa Dispholidus Droniophis Drymarchon Drymobitis Dryophis Elaphe Elapomorphus Ehpops Langebartel 1968 Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Walter 1887 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Morgans & Heidt 1978 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Walter 1887 Langebartel 1968 Lubosch 1933 Albright & Nelson 1959 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Auen & Langebartel 1977 September 1982 Tanner, Avery: Buccal Floor of Reptiles 285 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Enhydrus Langebartel 1968 Langebartel 1968 Langebartel 1968 Enulius Langebartel 1968 Farancia Langebartel 1968 Ficimia Langebartel 1968 Fimbrios Langebartel 1968 Langebartel 1968 Langebartel 1968 Geophis Langebartel 1968 Haldea Langebartel 1968 Haplopeltura Langebartel 1968 Langebartel 1968 Langebartel 1968 Heterodon Weaver 1965 Langebartel 1968 Langebartel 1968 Langebartel 1968 Homalopsis Langebartel 1968 Lampmpeltis Langebartel 1968 Leptodeira Langebartel 1968 Leptophis Langebartel 1968 Manolepis Langebartel 1968 Masticophis Langebartel 1968 Mehelya Langebartel 1968 Langebartel 1968 Langebartel 1968 Matrix Sondhi 1958 Sondhi 1958 Sondhi 1958 Sondhi 1958 Nerodia Langebartel 1968 Oldham, Smith & Miller 1970 Morgans & Heidt 1978 Varkey 1979 Langebartel 1968 Oldham, Smith & Miller 1970 Varkey 1979 Langebartel 1968 Varkey 1979 Ninia Langebartel 1968 Nothopsis Langebartel 1968 Langebartel 1968 Langebartel 1968 Opheodrys Langebartel 1968 Cundall 1974 Cundall 1974 Oxyhelis Langebartel 1968 Oxyrhahdinium Langebartel 1968 Pitiiophis Smith & Warner 1948 Bullock & Tanner 1966 Langebartel 1968 Oldham, Smith & Miller 1970 Oldham, Smith & Miller 1970 Psamnodynastes Langebartel 1968 Rhadineae Langebartel 1968 Rhadinelia Langebartel 1968 Rliinocheilus Langebartel 1968 Salvadora Langebartel 1968 Sibynomorphus Langebartel 1968 Langebartel 1968 Langebartel 1968 Sonora Langebartel 1968 Tantilla Langebartel 1968 286 Great Basin Naturalist Vol. 42, No. 3 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Thamnophis Toluca Trimorphodon Tropidonotus Xenodermus Xenodon Elapidae Acanthophis Aspidelaps Bungarus Calliophis Demansia Dendrospis Denisonia Doliophis Elaps Elapsoidea Furina Hemachatus Hemtbungarus Leptomicnirus Maticora Micruroides Micrurus Naja Notechis Ogmodon Pseudechis Pseudelaps Ultocalamus Bullock & Tanner 1966 Langebartel 1968 Oldham, Smith & Miller 1970 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Weaver 1965 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Smith & Warner 1948 Langebartel 1968 Langebartel 1968 Kanial, Hamouda & Mokhtar 1970 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Sewertzoff 1929 Langebartel 1968 Oldham, Smith & Miller 1970 Lubosch 1933 Langebartel 1968 Anthony & Serra 1949 Langebartel 1968 Langebartel 1968 Langebartel 1968 Lubosch 1933 Langebartel 1968 Langebartel 1968 Kesteven 1944 Langebartel 1968 Aven & Langebartel 1977 Lubosch 1933 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Langebartel 1968 Kesteven 1944 September 1982 Tanner, Avery: Buccal Floor of Reptiles 287 Table 1 continued. Genus Hyoid Tongue Musculatupe Nerves Hydrophidae Aipysurus Langebartel 1968 Langebartel 1968 Langebartel 1968 Hydrophis Langebartel 1968 Langebartel 1968 Langebartel 1968 Kerilia Langebartel 1968 Lapemis Smith & Warner 1948 Langebartel 1968 Laticauda Langebartel 1968 Langebartel 1968 Langebartel 1968 Pelamis Langebartel 1968 Langebartel 1968 Langebartel 1968 Thalasophina Langebartel 1968 Viperidae Aspis Langebartel 1968 Langebartel 1968 Langebartel 1968 Atheris Langebartel 1968 Atractaspis Langebartel 1968 Langebartel 1968 Langebartel 1968 Bitis Langebartel 1968 Causus Langebartel 1968 Haas 1952 Langebartel 1968 Langebartel 1968 Cerastes Langebartel 1968 Langebartel 1968 Langebartel 1968 Echis Langebartel 1968 Langebartel 1968 Langebartel 1968 Pseudocerastes Langebartel 1968 Vipera Furbringer 1922 Edgeworth 1935 Langebartel 1968 Langebartel 1968 Langebartel 1968 Crotalidae Agkistrodon Smith & Warner Langebartel 1968 Langebartel 1968 1948 Kardong 1973 Langebartel 1968 Bothrops Langebartel 1968 Langebartel 1968 Langebartel 1968 Crotalus Langebartel 1968 Langebartel 1968 Langebartel 1968 Oldham, Smith & Oldham, Smith & Miller 1970 Miller 1970 Lachesis Langebartel 1968 Lubosch 1933 Lubosch 1933 Langebartel 1968 Langebartel 1968 Sistntrus Langebartel 1968 Trimeresurus Langebartel 1968 Order Crocodilia Crocodylidae Alligator Reese 1915 Sewertzoff 1929 Reese 1915 Reese 1915 Furbringer 1922 Lubosch 1933 Lubosch 1933 Edgeworth 1935 Edgeworth 1935 Poglayen-Neuwall Chiason 1962 Chiason 1962 Poglayen-Neuwall 1953b 1953b Caiman Schumacher 1973 Schumacher 1973 Schumacher 1973 288 Great Basin Naturalist Vol. 42, No. 3 Table 1 continued. Genus Hyoid Tongue Musculature Nerves Crocodylus Furbringer 1922 Sondhi 1958 Frank & Smit 1974 Sewertzoff 1929 Sondhi 1958 Camp 1923 Edgeworth 1935 Kesteven 1944 Sondhi 1958 Poglayen-Neuwall 1953b Kesteven 1944 Poglayen-Neuwall 1953b Cavialidae Gavialis Furbringer 1922 Sondhi 1958 Sondhi 1958 Sondhi 1958 Sondhi 1958 attempts have not led to a completely satis- factory understanding of the hyoid deriva- tives, and the homologies of the hyoid con- stituents cannot be ascertained without comparative embryological information on the development of the reptiles' buccal floor. Thus, our interpretation of the hyoids of rep- tiles should be considered tentative at best. The tentativeness of our present under- standing of the homologies of many of these structures, when considered for the reptiles as a whole, is indicated in a recent study by Frank and Smit (1974). They consider the early ontogeny of the columella auris of Crocodylus niloticus, and its relationship to the reptilian hyoid arch is reviewed and dis- cussed in detail. Considerable effort is ex- pended in the clarification of terms and in the description of particular morphological structures that have been in some confusion as a result of misunderstandings. Frank and Smit (1974) were trying to establish a model to be used, and to stimulate subsequent re- search. We hope that such studies will be un- dertaken and will clarify structural homo- logies for each of the remaining arches occurring in reptiles. Ontogenetic studies leading to an understanding not only of rep- tiles in general but also of each reptilian or- der are needed before we can consistently as- sign exact anatomical limits to structures and provide an appropriate name for each mor- phological entity. Until such are available, however, we must remain within the limits of our present knowledge. The hyoid apparatus of reptiles has been described by Romer (1956) as having a me- dian ventral piece, the copula, which forms the body of the hyoid (= corpus hyoidem BH). Extending anteriorly from the corpus hyoideimi is a medial process, the processus lingualis ( = processus entoglossus PL), which terminates in the substance of the tongue. Laterally the corpus gives off three paired horns or comua that extend posterodorsally around the throat, ending freely or attached to the stapedial structures. The anteriormost pair of comua are the hyoid comua (HC), which, at their distal ends, articulate with the most lateral of the hyoid bars or epihyals (EH). The second pair of comua are elon- gated bars (usually cartilagenous) forming the first ceratobranchials (CBI) and attaching dis- tally to the epihyals by way of short cartila- ginous bars on each side of the first epibran- chials (EBI). The third and last processes of the corpus hyoideum are a pair of posteriorly extending rods, the second epibranchials (EBII), a pair of short cartilaginous bars that articulate by their distal ends with the epi- branchials of the first and second arches (Figs. 1-3). The association of the hyoid with, and its attachment to, the skull is of evolutionary im- portance. In primitive lizards, such as some Gekkonidae, the hyoid is attached to the ex- tracolumella. Thus the hyoid apparatus main- tains its association with the stapedial struc- tures that may be derived from its dorsal extremities. This point of union is or is not persistent as indicated by Versluys (1898 and 1904) and, in some individuals, is transferred to the paraoccipital process of the opisthotic bone. The possession of all three comua is con- sidered the primitive condition and is demon- strated in Sphenodon, where all the comua are long, slender, and well-developed struc- tures. A few gekkos and xantusiids have an incomplete third arch with a slight break be- tween ceratobranchial II and epibranchial II. In the Iguanidae (Avery and Tanner 1971) all September 1982 Tanner, Avery: Buccal Floor of Reptiles 289 three arches are present, but lack their distal connections in some cases. Rhynchocephalia The hyoid of Sphenodon has been discussed by Osawa (1898, Howes and Swinnerton 1901, Furbringer 1922, Edgeworth 1931, 1935, and Rieppel 1978. The hyoid apparatus of Sphenodon (Fig. 1) is simple with all elements present. The basi- hyoid is broad with a short lingual process ex- tending anteriorly. Laterally the basihyoid extends as projections corresponding to the hyoid cornua but not distinctly separate from the basihyoid. At their distal ends, the cornua articulate with epihyals that extend straight posteriorly. The basihyoid also has a pair of posterior projections, the second ceratobran- chials, that are widely separated and curve laterally at their distal ends. The first cerato- branchials articulate with the basihyoid later- al to the point of origin of the second cerato- branchials. They curve and closely approach the distal ends of the epihyals. Rieppel (1978) illustrated the hyoid apparatus and its associ- ated muscles. A taxonomic survey provides a general overview of this order: Chelonia The hyoid apparatus of turtles has been de- scribed by the following: Chelidae Chelodina (Furbringer 1922) Dermatemydidae Dermatemys (Furbringer (1922) Chelydridae Cheydra (Furbringer 1922, Edgeworth 1935, Schumacher 1973), Kinosternon (Furbringer 1922, Schumacher 1973), Ster- notherus (Furbringer 1922, Schumacher J973), Chrysemys (Furbringer 1922, Ash- ley 1955, Schumacher 1973), Cuora (Fur- bringer 1922), Clemmys (Siebenrock 1898, Furbringer 1922, Schumacher 1973), Emys (Walter 1887, Furbringer 1922, Schuma- cher 1973), Geochelone (Bojanus 1819, Furbringer 1922, Edgeworth 1935, Schumacher 1973), Terrapene (Furbringer 1922). Trionychidae Lissemys (Furbringer 1922, Sondhi 1958, Schumacher 1973), Trionyx (Siebenrock 1898, Sondhi 1958, Furbringer 1922, Schumacher 1973). Cheloniidae Caretta (Furbringer 1922, Schumacher 1973). Dermochelyidae Dermochelys (Schumacher 1973). Schumacher (1973) has treated the hyoids of turtles and crocodilians extensively in this series, so our discussion will serve as a gener- al review. The hyoid apparatus of turtles has been de- scribed briefly by Bojanus (1819) and figured by Mitchell and Morehouse (1863). More Fig. 1. Hyoid apparatus of Sphenodon punctatum (USUN 029429): BH-body of hyoid, (basihyoid) CBl-first ceratobranchial, CBll-second ceratobranchial, EBl-first epibranchial, EBll-second epibranchial, EH-epihyal, HC-hyoid comu, PL-processus lingualis. 290 Great Basin Naturalist Vol. 42, No. 3 Fig. 2.— Hyoid apparatus of A, Chelydra serpentina (Southern Connecticut State College, 598), ventral view; B, Caiynan sclerops, ventral view; C, Caiman sclerops, lateral view (SCSC 585). complete reports include those of Siebenrock (1898), Furbringer (1922), Versluys (1936), Gnanamuthu (1937), and Sondhi (1958). The hyoid is more ossified than that of most liz- ards and snakes. In Trionyx and Lissemys the hyoid has a body with a lingual process equipped with a hypoglossum (Sondhi 1958); this is a leaflike plate of cartilage loosely attached to its ven- tral side. The hyoid comua are greatly re- duced and form knoblike projections from the body. The second ceratobranchials extend posteriorly from the body as subcylindrical structures. The body of the hyoid is composed of three pairs of serially arranged cartilaginous blocks. The most anterior part has on its lat- eral margins very short anterior projections. The middle pair of plates bear the articu- lating surfaces for the hyoid comua. The pos- terior pair of plates are completely fused to the middle pair and have between them and the middle plates a diamond-shaped inter- space. Posteriorly the last pair of plates pro- vides facets for the articulations of the second ceratobranchials. In Chelydra the hyoid is more solidly constructed, consisting of bone except for its anterior end, the ceratohyals, and the epihyals, which are cartilage (Fig. 2 A). The possession of a hypoglossum by turtles appears to be unique. The structure was first described by Stannius (1856) as an entoglos- sum. The term hypoglossum was first used by Furbringer (1922), who described it as the part not entering the tongue. Nick (1913) and Versluys (1936) observed that in turtles, with the exception of Dermochelys, the hypoglos- sum is platelike, unpaired, and lies ventral to the lingual process. Nick (1913) also suggests that the hypoglossum is a chondrification of connective tissue of the tendinous plate. The hypoglossum is extensive in Trionyx, in which it may have two slender posterior strips or be an elongate plate, rounded at each end and extending anteriorly from the middle com- ponents of the body of the hyoid almost to the symphysis of the mandible. Sondhi (1958) suggested that the hypoglossum functions to raise or lower the buccal floor by means of two muscles (Mm. entoglosso-hypoglossalis and hypoglosso-lateralis) attached to its dor- sal surface and extending to the processus en- toglossus and the buccal floor. In other gen- era, Chelydra, Chrysemys, Pseudemys, and Sternotherus, it is proportionally smaller and varies in shape (Fig. 3). Hacker and Schu- macher (1955) figure it for Testudo and de- scribe the M. entoglosso-glossus that serves as an attachment between the hypoglossum and the processus lingualis. In Gopherus agassizi, the hypoglossum is elongate and slender with a median ridge ventrally and a convexity dor- sally. It is closely associated with the process- us lingualis. A paired muscle (M. entoglosso- glossus) is attached to its dorsal surface on September 1982 Tanner, Avery: Buccal Floor of Reptiles 291 Fig. 3. The hypoglossum of five genera of turtles, ventral views: A, Chrysemys picta (SCSC 602); B, Sternotherus odoratus (SCSC 476); C, Pseudemys scripta (BYU 40343); D, Chelydra serpentina (BYU 33642); E, Trionyx spinifera (SCSC 596); F, Gopherus agassizi (BYU 30084). each side, just lateral to the median con- cavity. These muscles extend dorsally and an- teriorly to insert in the connective tissues sur- rounding the processus lingualis. The pointed anterior end of the hypoglossum extends beyond the end of the processus lingualis. In the few examples we have seen, the hy- poglossum of terrestrial tortoises appears to be more slender and with better developed muscular attachments to the hyoid apparatus than in other turtles. The hyoid comua are short cartilaginous knobs covered ventrally by the M. cerato- hyoideum. The first ceratobranchials are long, subcylindrical, rodlike bones that artic- ulate with the middle component of the body of the hyoid. They extend posteriorly to curve dorsally and partially surround the neck, where they lie embedded in the M. ceratohyoideus. The second ceratobranchials are composed of proximal bony parts and distal cartila- ginous parts. The distal ends girdle the poste- rior part of the neck and lie beneath the M. omohyoideum. A ligament connects the base of each second ceratobranchial with the ante- rior part of each hyoid comu. Crocodilia In Alligator, Crocodylus, and Gavialis the hyoid apparatus consists of the body of the hyoid and a pair of posterior projections. The hyoid comua and all other processes are ab- sent. Sondhi (1958) has described the struc- tures in Gavialis in detail. The body of the hyoid is the most prominent part of the appa- ratus and forms an inverted triangular car- tilaginous plate. There is a deep notch in the posterior margin, and laterally it bears a fac- et for the articulation of the posterior projec- tion. The hyoid lies dorsal to the M. mylo- hyoideus, ventral to the glottis, and anterior to part of the trachea. Anteriorly the body is covered in part by the Mm. hyoglossus and genioglossus. The posterior projections are rodlike, cartilaginous, and extend post- eromedially, gradually becoming flattened, compressed, and twisted. A ligament con- nects these projections with fused rodlike structures closely adhering to the post- erolateral borders of the body and probably corresponding to the second ceratobranchials of other reptiles. The above description of Gavialis corre- sponds to our findings in Caiman except that the body of the hyoid of the latter is not triangular, but broadly rectangular and, from a dorsal view, similar to a wide-bladed shovel (Fig. 2-B,C). There is a shallow notch pos- teriorly, and the posterior projections are bone proximally and expand into flat sheets of cartilage distally. We did not find a liga- ment extending dorsolaterally onto the cer- vical area from the ends of the posterior projections. Lacertilia The hyoid of lizards has been examined by the following: Gekkonidae Cnemaspis (Richter 1933), Coleonyx (Camp 1923, Kluge 1962), Eublepharis 292 Great Basin Naturalist Vol. 42, No. 3 Table 2. Publications, not previously cited, dealing with topics peripheral to the buccal floor. A. Osteology 1. Chelonia Ashley 1955, Chelydra, Chrysemys Goppert 1903, Testudo 2. Rhynchocephalia Goppert 1900, Sphenodon Lakjer 1927, Sphenodon Rieppel 1979, 1981, Sphenodon 3. Lacertila Barrows and Smith 1947, Xenosaunis Beddard 1905a, Ihomastix Bellairs 1950, Anniella Criley 1968, Barisia, Elgaria, Gerrhonotus Duda 1965, Agama El Toubi 1938, Scincus El Toubi 1947a, Agama El Toubi 1947b, UromasHx El Toubi and Kamal 1959a, Chalcides El Toubi and Kamal 1959b, Chalcides Elyal-Giladi 1964, Agama, Chalcides George 1954, UromasHx Goppert 1903, Amphisbaena, Calotes, Cnemido- phorus, Lacerta, Mabuya, Platydactylus Iyer 1942, Calotes Iyer 1943, Calotes Kingman 1932, Eumeces Lakjer 1927, Ameiva, Anguis, Amphisbaena, Ca- lotes, Chalcides, Chamaelo, Cordylus, Eumeces, Gekko, Hyperodapedon, Heloderma, Iguana, Lialis, Lygosoma, Phrynosoma, Pygopus, Lacer- ta, Tiligua, Trogonophis, Uromastix, Varanus Mahendra 1949, Hemidactylus Malam 1941, Gerrhosaurus Norris and Lowe 1951, Phrynosoma Parker 1880, Lacerta, Agama Rathor 1969, Ophiomorus Rice 1920, Eumeces Siebenroek 1892a, Uroplatus Siebenrock 1892b, Scincus Siebenroek 1893, Brooksesia Siebenrock 1894, Lacerta Siebenrock 1895, Agama Sinitsin 1928, Alopogloscus, Ameiva, Anadia, Bach- ia, Callopistes, Cercosaura, Centropyx, Cnemido- phorus, Dracaena, Dicrodon, Echinosaura, Ec- leopus, Euspondylus, Gymnophthalmus, Iphisa, Leposoma, Neusticurus, Ophiognomon, Pan- todactylus, Prionodactylus, Pholidobolus, Pructo- porus, Scolecosaurus, Teius, Tretioscincus, Tupinambis Tilak 1964a, Uromastix Toerien 1950, Anniella Webb 1951, Oedura, Palmatogecko Weiner and Smith 1965, Crotaphytus Young 1942, Xantusia Zangerl 1944, Amphisbaena, Bipes, Geocalamus, Leptosternon, Monopelitis, Rhineura, Trogonophis Table 2 continued. 4. Ophidia Herman 1961, Echis, Vipera Herman 1965, Calamaria Boltt and Ewer 1954, Bitis Dullemeijer 1956, Vipera Dullemeijer 1959, Bitis, Crotalus, Trimeresurus, Vipera Kardong 1974, 1977, Agkistrodon Liem, Mark and Rabb 1971, Azemiops Goppert 1903, Python, Tropidonotus McKay 1889, Acanthrophis Rosenberg 1968, Bungarus Varkey 1979, Nerodia 5. Crocodilia Chiasson 1962, Alligator Goppert 1903, Crocodylus B. Myology 1. Chelonia Adams 1919, Chelydra Ashley 1955, Chelydra, Chrysemys Schumacher 1956, Amyda, Chelodina, Chelonia, Caretta, Clemmys, Dogania, Emydura, Emys, Eretmochelys, Graptemys, Hardella, Macro- chelys, Hydromedusa, Pelomedusa, Pelusios, Pla- tysternon, Podocnemis, Testudo, Trionyx Shah 1963, Chelodina, Deirochelys 2. Rhynchocephalia Adams 1919, Sphenodon Rieppel 1978, Sphenodon 3. Lacertila Adams 1919, Iguana, Varanus Bradley 1903, Agama, Gekko, Lacerta, Pseudopus, Varanus Brock 1938, Gymnodactylus Davis 1934, Crotaphytus George 1948, Uromastix lordansky 1970, Agama, Cordylus, Eumeces, Gek- ko, Lacerta, Ophiosaurus, Teratoscincus, Varanus Norris and Lowe 1951, Phrynosoma Rathor 1969, Ophiomorus Tornier 1904, Chamaeleo 4. Ophidia Adams 1925, Natrix Bergman 1961, Echis, Vipera Bergman 1965, Calamaria Boltt and Ewer 1954, Bitis Cowan and Hick 1951, Thamnophis Dullemeijer 1956, Vipera Dullemeijer 1959, Bitis, Crotalus, Trimeresurus, Vipera Haas 1930, Amblycephalus, Calharia, Calamaria, Cylindrophis, Eryx, Ilysia, Oxybelis, Silybura, Xenopeltis Haas 1931a, Acrochordus, Amblycephalus, Atrac- taspis, Atractus, Bungarus, Calabaria, Cala- September 1982 Tanner, Avery: Buccal Floor of Reptiles 293 Table 2 continued. maria, Causus, Cerberus, Chersydrus, Cylin- drophis, Dasypeltis, Dispsadomonphus, Elaps, Eryx, Glauconia, Ilysia, Lachesis, Leptognathus, Naja, Oxybelis, Pelamis, Python, Poly- odontophis, Silyura, Typhlops, Xenodon, Xenopeltis Haas 1931b, Acrodordiis, Atractaspis, Causus, Cer- berus, Chersydrus, Cylindrophis, Dasypeltis, Dispholidus, Leptognathus, Petalognathus, Poly- odontophis, Scaphiophis, Xenodon Haas 1952, Causus Heymans 1970, Matrix Heymans 1975, Aparallactus, Atractaspis, Chilorhinophis Kochva 1958a, Vipera Kochva 1958b, Agkistrodon, Aspis, Atheris, Atrac- taspis, Bitis, Bothrops, Causus, Crotalus, Echis, Natrix, Naja, Ophiophagus, Pseudocerastes, Vi- pera, Walterinnesia Kardong 1974, Agkistrodon Liem, Mark, and Rabb 1971, Azeniiops McKey 1889, Acanthrophis Rosenberg 1968, Bungarus Rosenberg and Cans 1976, Elachistodon Crocodilia Adams 1919, Alligator Chiasson 1962, Alligator C. Miscellaneous 1. Chelonia Johnson 1922, Branchial pouch derivatives, Che- lydra, Chrysemys Goppert 1900, Larynx, Chelonia, Dermochelys, Emtjs, Testudo Siebenrock 1900, Larynx, Testudo 2. Lacertila Goppert 1900, Larynx, Amphishaena, Platydac- tylus, Tiliqiia Perrier 1902, Thymus and thyroid glands, Lacerta Saint-Remy and Prenant 1904, Thymus and thy- roid glands, Anguli, Lacerta Sidkey 1967, Carotid Sinus, Chalcides, Scincus 3. Ophidia Goppert 1900, Larynx, Coronella, Python, Tropidonotus Kroll 1973, Taste buds, Leptotyphlops Saint-Remy and Prenant 1904, Thymus and thy- roid glands. Coluber, Tropidonotus Van Bourgondien and Bother 1969, Cephalic arte- rial patterns, Agkistrodon, Crotalus. Lachesis, Slstrurus 4. Crocodilia Goppert 1900, Larynx, Crocodylus Siebenrock 1899, Larynx, Crocodylus (Cope 1892, Camp 1923), Gekko (Camp 1923, Richter 1933), Gehydra (Richter 1933), Gymnodactylus (Richter 1933), Hemidactylus (Zavattari 1908, Richter 1933, Edgeworth 1935), Phyllodactylus (Cope 1892), Ptychozoon (Richter 1933), Tarentola (Richter 1933), Uroplatus (Ver- sluys 1898, 1904, Camp 1923, Edgeworth 1935). Dibamidae Dibamus (Rieppel 1981). Iguanidae Amblyrhynchus (Avery & Tanner 1971), Anolis (Cope 1892), Basiliscus (Zavattari 1908), Brachylophus (Camp 1923, Avery & Tanner 1971), Callisaurus (Cox & Tan- ner 1977), Chalarodon (Avery & Tanner 1971), Chamaeleolis (Beddard 1907), Con- olophus (Avery & Tanner 1971), Copho- saurus (Cox & Tanner 1977), Crotaphytus (Cope 1892, Robison & Tanner 1962), Ctenosaura (Oelrich 1956, Avery & Tan- ner 1971), Cyclura (Avery & Tanner 1971), Dipsosaurus (Cope 1892, Avery & Tanner 1971), Enyaliosaurus (Avery & Tanner 1971), Holbrookia (Cox & Tanner 1977), Iguana (Edgeworth 1935, Avery & Tanner 1971, Oldham & Smith 1945), Ophirus (Avery & Tanner 1971), Phryno- soma (Cope 1892, Camp 1923, Richter 1933, Jenkins & Tanner 1968), Polychrus (Richter 1933), Sauromalus (Avery & Tan- ner 1964, 1971), Sceloporus (Cope 1892), Tropidurus (Zavattari 1908, Edgeworth 1935), Uma (Cox & Tanner 1977), Uro- saurtis (Fanghella, Avery & Tanner 1975), Uta (Fanghella, Avery & Tanner 1975). Agamidae Agama (Edgeworth 1935, El Toubi 1947, Harris 1963, Eyal-Giladi 1964), Amphibo- luriis (Richter 1933), Calotes (Zavattari 1908, Camp 1923, Richter 1933, Edge- worth 1935, Iyer 1943), Ceratophura (Richter 1933), Chlamydosaurus (Beddard 1905), Cophotis (Richter 1933), Draco (Richter 1933), Hydrosaurus (Richter 1933), Leiolepis (Richter 1933), Lyr- iocephaltis (Richter 1933), Otocryptis (Richter 1933), Phrynocephahis (Richter 1933, Kesteven 1944), Physignathus (Kes- 294 Great Basin Naturalist Vol. 42, No. 3 Fig. 4. Hyoid apparatus of Tarentola annularis (BYU 18123): A, ventral view; B, lateral view. teven 1944), Uromastix (Islam 1955, Tilak 1964a,b). Chamaeleonidae Chamaeleo (Zavattari 1908, Edgeworth 1935, Gnanamuthu 1937, Jollie 1960). Scincidae Acontias (Rieppel 1981), Acontophiops (Rieppel 1981), Chalcides (Richter 1933, El Toubi 1938, El Toubi & Kamal 1959a,b), Eumeces (Cope 1892, Zavattari 1908, Richter 1933, Nash & Tanner 1970), Lygosoma (Richter 1933), Mahuya (Rich- ter 1933, Gnanamuthu 1937, Rao & Ra- maswami 1952), Nessia (Richter 1933), Riopa (Richter 1933), Tiliqua (Beddard 1907), Scincus (Richter 1933), Trachy- saurus (Beddard 1907), Typhlosaurus (Rieppel 1981). Cordylidae Cordylus (Beddard 1907, Camp 1923, Richter 1933, Edgeworth 1935), Gerrho- saurus (Camp 1923), Zonurus (Camp 1923). Lacertidae Acanthodactylus (Richter 1933), Lacerta (Walter 1887, Zavattari 1908, Richter 1933, Edgeworth 1935), Ophisops (Richter 1933). Teiidae Ameiva (Richter 1933, Fisher & Tanner 1970), Cnemidophorus (Cope 1892, Fisher & Tanner 1970), Neusticurus (Richter 1933), Tupinambis (Zavattari 1908, Reese 1932, Edgeworth 1935, Jollie 1960). Anguinidae Anguis (Richter 1933), Gerrhonotus (Wal- ter 1887, Cope 1892), Ophiosaurus (Wal- ter 1887). Xenosauridae Shinosaurus (McDowell & Bogert 1954), Xenosaurus (McDowell & Bogert 1954, McDowell 1972). September 1982 Tanner, Avery: Buccal Floor of Reptiles 295 Fig. 5. Hyoid apparatus of Coleonyx variegatus (BYU 18796): A, ventral view; B, lateral view. Helodermatidae Heloderma (Cope 1892, McDowell & Bo- gert 1954). Varanidae Varanus (Richter 1933, McDowell & Bo- gert 1954, Sondhi 1958). Lanthanotidae Lanthanotus (McDowell & Bogert 1954, Rieppel 1981). Anniellidae Anniella (Cope 1892, Rieppel 1981). Anphisbaenidae Amphisbaena (Camp 1923, Richter 1933, Jollie 1960), Monopeltis (Richter 1933), Rhineura (Cope 1892). Xantusidae Xantusia (Cope 1892, Savage 1963). Most lizards have a hyoid consisting of a basihyal (corpus hyoideum) with a pair, each, of anterior and posterior comua as described by Cope (1892), Zavattari (1908), Furbringer (1922), Camp (1923), Versluys (1936), DeBeer (1937), Gnanamuthu (1937), Mahendra (1947), Rao and Ramaswami (1952), McDowell and Bogert (1954), Oelrich (1956), Romer (1956), Sondhi (1958), Jollie (I960), Robison and Tanner (1962), Avery and Tan- ner (1964), Jenkins and Tanner (1968), Fisher and Tanner (1970), Nash and Tanner (1970), Avery and Tanner (1971), Rieppel (1981), and others. For the remainder of this dis- cussion we will use the hyoid nomenclature followed by Romer (1956) as described ear- lier. The hyoids of the geckos Coleonyx, Gek- ko, Aristelliger, Hemidactylus, Phyllodactylus, Thecadactylus, and Eublepharis have been described, and we figure Tarentola (Fig. 4) and Coleonyx (Fig. 5). In most, the body of the hyoid is small and slender, with a long rodlike lingual process extending anteriorly. A pair of hyoid comua extend laterally; in 296 Great Basin Naturalist Vol. 42, No. 3 Fig. 6. Hyoid apparatus, ventral views: A, Brachylophus brevicephahts (BYU 32663); B, Sattromalus obesus (BYU 21728). some species these form sigmoid curves, and in others they are straight rods. Articulating with the distal extremes of the hyoid comua are the epihyals. Extending posteriorly from the body of the hyoid as a pair of short or long rods are the second ceratobranchials. A third set of arches, the first ceratobranchials, articulate at the point of attachment between the hyoid comua and the body. The basic pattern is retained throughout the Gekkota, with some variation in the shape of the hyoid comua; also, the first ceratobranchials, epi- hyals, or both may be lost in some genera. In the Dibamidae, Rieppel (1981) has de- scribed the hyoid of Dibamus as having a posteriorly bifurcated basihyal with an elon- gated entoglossal process. The bony first ce- ratobranchials that articulate with the post- erolateral limbs of the basihyal are shorter in Dibamus as compared to Anniella. He in- dicates a major specialization exists in that there are a pair of cartilaginous rods that support the aditus laryngis and approach but do not fuse to the posterolateral limbs of the basihyal. These he considers to be hypohyals (hyoid comua of Romer). The hyoids of the iguanine lizards Ambly- rhynchus, Brachylophus, Conolophus, Cteno- saura, Cyclura, Dipsosaurus, Iguana, and Sauromalus and Malagashe iguanids Chalaro- don and Oplurus have been investigated by Avery and Tanner (1971). Because these liz- ards possess all three arches of the hyoid ap- paratus, they are considered primitive (Fig. 6-A). The body of the hyoid (basihyal) is triangular in all the above genera except Oplurus and Sauromalus, in which it forms a broad flattened sheet of cartilage. In all the genera the hyoid comu (hypohyal) is short and stout; it extends out from the body of the hyoid at right angles or projects slightly ante- rior to the body. Posterior to the body, the second ceratobranchials extend along the trachea and, in all genera except Oplurus and Sauromalus, lie close together. In the latter two genera the second ceratobranchials are widely separated by the bulk of the trachea (Fig. 6-B). In none of the genera are the sec- September 1982 Tanner, Avery: Buccal Floor of Reptiles 297 Fig. 7. Hyoid apparatus, ventral views: A, Sceloportis magister (BYU 30310); B, Holbrookia maculata (BYU 15752); C, Phrynosoina platyrbinos (BYU 22830). end ceratobranchials attached distally to the other arches. In some genera, particularly Iguana, the distal extremes of these processes attach to the skin and provide support for movement of the dewlap. The first ceratobranchials articulate prox- imally w^ith the body of the hyoid between the origins of the second ceratobranchials and the hyoid comua. They are elongated, thin rods that taper to points distally and curve dorsolaterally to the sides of the neck, where they articulate with the epihyals (cer- tohyals). The epihyals articulate between the hyoid comua and the first ceratobranchials and form the most lateral extensions of the hyoid apparatus. At their proximal ends the epihyals are expanded into bladelike process- es that extend medially toward the hyoid body. These processes are not developed to any degree in Chalarodon and Opiums. Among the other iguanids studied and de- scribed by one of us are the hyoids of Crota- phytus, Holbrookia, Phnjnosonia, and Uta. We figure Sceloporus magister and Hol- brookia maculata (Figs. 7-A & B) as represen- tatives of the sceloporine genera. The basic pattern described in the iguanines is main- tained with the following exceptions. In Phrynosoma the second ceratobranchials are greatly reduced, and the first ceratobran- chials and epihyals are noticeably thickened (Fig. 7-C); the basihyoid is a laterally extend- ed plate. Anolis has an exceptionally elon- gated hyoid apparatus, with the second ce- ratobranchials extending posteriorly along the midline forming approximately two- thirds the length of the entire hyoid appa- ratus. This anatomical development is associ- ated with the functional dewlap (Fig. 8). In the agamids, the following were exam- ined: Agama (Duda 1965, Hass 1973), and Figure 9; Calotes, Draco, and Sitana (Gnana- 298 Great Basin Naturalist Vol. 42, No. 3 CBl CBll muthu 1937), Chlamydosaurus (Beddard 1905, DeVis 1883), Phrynocephalus (Haas 1973), Physignathus (Kesteven 1944), and Uromastyx (Poglayen-Neuwall 1954, Versluys 1898, El Toubi 1947b, Tilak 1964b). In gen- eral, the agamid hyoids resemble closely those of the iguanids. In Uromastyx the basi- hyoid is slender and laterally extended; the hyoid comua are directed anterolaterally (Ti- lak 1964b). The short and widely separated second ceratobranchials extend posteriorly from the basihyoid. The first ceratobranchials extend posteriorly from the basihyoid. The first ceratobranchials articulate at the union of the hyoid comua and the basihyoid. They comprise the longest elements of the hyoid. The epihyals attach to the distal ends of the hyoid comua and have, at their distal ends, epibranchials that may attach to the distal end of the first ceratobranchials. In Agama (Fig. 9) the hyoid is similar except that the basihyoid is more massive and the second ce- ratobranchials are aligned more closely to- gether. In Calotes and Draco the hyoids are elongated and narrow. The second cerato- branchials are exceptionally long and slender, lying close together at the midline, whereas CBll Fig. 8. Hyoid apparatus, ventral view: Anolis caroli- nensis (BYU 13768). Fig. 9. Hyoid apparatus of Agama agama (BYU 18147), ventral view. September 1982 Tanner, Avery: Buccal Floor of Reptiles 299 Fig. 10. Hyoid apparatus: A, Chamaeleon namagyensis (USNM 161275); B, Chamaeleon brevicornis (BYU 12422), ventral views; C, same as B, lateral view. the epihyals are very short and not connected by epibranchials. In Chlamydosaurus the basihyoid is massive and bears two homUke projections; these extend laterally to articu- late with the hyoid comua, which form short tapering tips on these projections. The sec- ond ceratobranchials appear to have been lost unless they are represented by two very small knobs on the posteromedial border of the basihyoid. The first ceratobranchials are extremely elongated, extending post- erolaterally and composed of two pieces. The very long proximal piece articulates distally with the second piece, which is about one- fifth the length of the proximal. The epihyals are short or slender, and articulate at the point where the hyoid comua and the lateral projections of the basihyoid attach. In Phys- ignathus the hyoid exhibits a normal struc- ture except that the first ceratobranchials are much longer than the second cerato- branchials. In Chamaeleo the hyoid is distinctly differ- ent, with the basihyoid being little more than the basal part of the lingual process. The hyoid comua extend anterolaterally about a third the length of the lingual process. The first ceratobranchials extend laterally and are short. The epihyals are small and attach to the hyoid cornua about half the distance from their distal ends. The second cerato- branchials are lost (Fig. 10-A, B, and C). Gnanamuthu (1937) described the hyoid ap- paratus for Chamaeleo carcaratus and re- viewed previous studies of its function. In the Scincidae the hyoids of Scincus (El Toubi 1938), Eumeces (Nash and Tanner 1970), Fig. 11, Mabuya (Richter 1933), and 300 Great Basin Naturalist Vol. 42, No. 3 EBI Fig. 11. Hyoid apparatus of Eumeces gilberti (BYU 31956), dorsal view. (After Nash and Tanner 1970) Chalcides (Furbringer 1922, Richter 1933) have been described. All three arches are present and assume an unspecialized pattern. In all the basihyoid is broad rather than nar- row, and the second ceratobranchials are very short and widely separated. The first ce- ratobranchials are elongate and slim. The hyoid comua are short and slim, and articu- late distally with the epihyals, which vary in form. They are simple rods in Eumeces and have enlarged proximal ends in the remain- ing genera. In Scincus the enlarged ends are simple and spoonshaped, but in Chalcides and Mahuya the shape is complex. In both genera the enlarged end has a short flange ex- tending posterolaterally from the middle of the epihyal where the enlarged end termi- nates. These genera have a large hooklike second epibranchial associated with the distal end of the epihyal. It is attached in Chalcides and Scincus but separate in Eumeces and Mahuya. In all genera there is a short first epibranchial attached to the terminal end of the first ceratobranchial (Fig. 11). Rieppel (1981) has examined the limbless scincoid genera Acontias, Typhlosaurus, and Acontaphiops. Acontias is described as being like Anniella, with the basihyal having a slen- der entoglossal process and being bifurcated posteriorly with its distinct posterolateral limbs articulating with first ceratobranchials. Hypohyal processes (hyoid comua) are pres- ent in all species where they are T-shaped at their distal ends. In Typhlosaurus the hyoid is similar to Acontias, but the posterior first ce- ratobranchials are longer and hypohyals are absent. Rieppel calls attention to the fact that the hyoid of Typhlosaurus is identical to that of some Typhlopidae as described by List (1966) and Langebartel (1968). The hyoid of Acontophiops is similar to that of Typhlosaurus. In the teiid Tupinambis, the lingual pro- cess is detached from the basihyoid and em- bedded in the tongue. The second cerato- branchials are lost, and the epihyals and first ceratobranchials are connected by epibranchials. September 1982 Tanner, Avery: Buccal Floor of Reptiles 301 Fig. 12. Hyoid apparatus of Cnemidophorus tigris (BYU 31925). Dorsolateral view showing the detached lingual process (LP) and the extension of the body of the hyoid anteriorly as a spine. The lingual process is also detached in Cnemidophoms (Fig. 12). The hyoid extends anteriorly as a short spine similar to that of igiianids except for its smaller size. It is em- bedded in connective tissue ventral to the lingual process and the tongue. The hyoid comua extend anterolaterally from the basi- hyoid and articulate with the epihyals. The latter extend anteriorly, forming bladelike cartilages that serve as lateral supports for the posterior half of the tongue and extend laterally to lie adjacent to the mandible. The posterior part of the epihyal extends posteri- orly, curving laterally where it terminates as cartilage in loose connective tissue on the first ceratobranchial. Both ceratobranchials are present; the first extends posteriorly to terminate in the connective tissue with the cartilagenous first epibranchial. The epihyals and first ceratobranchials are not connected distally, although the ends are close together ip a common connective tissue. Ameiva lacks the second ceratobranchials. The first epibranchials are short, forming a knob on the end of the ceratobranchials. In both Ameiva and Cnemidophorus the detached lingual process extends anteriorly to approximately the forking of the tongue. Pos- teriorly it is tightly enclosed in connective tissue between the elongate M. hypoglossus. It terminates posteriorly, ventral to the lar- yngeal cartilages. In Angtiis (Anguidae) the hyoid is greatly reduced, with the second ceratobranchials and epihyals absent. The hyoid comua are enlarged and extend anteriorly to parallel the lingual process for most of its length. In Ger- rhonotus and Ophisaurus the second cerato- branchials are also lost. The epihyals are present, however, and articulate with the dis- tal ends of the hyoid comua, which are more laterally directed than in Anguis. In Varanus (Varanidae), the hyoid comua is complex and is composed of two articu- lating cartilaginous rods, called by Sondhi (1958:159-160) the portio proximalis (hyoid cornu) and the portio distalis (epihyal): Each has an anterior handlelike process and in life the two hooked ends cross each other beneath the tongue- sheath, with the handle of the portio proximahs lying dorsal to that of the portio distalis. According to Sondhi (1958:159-160), the proximal end of the portio proximalis fits into a roughly concave facet on the dorsolateral surface of the basihyoid, near the facet at which the posterior comua articulates. From this point the portio proximalis ex- tends obliquely upward, outward, and forward and at its termination curves inward to form the hook-shaped handle that is dorsoventrally flattened. The portio dis- talis is flattened at its proximal handlelike end, becomes rodlike as it passes backward and upward, and gradually 302 Great Basin Naturalist Vol. 42, No. 3 tapers at its distal end. It is disposed obliquely across the sides of the neck, its tapering end lying almost parallel to the proximal piece of the posterior cornua of its side. Sondhi also indicates that the portio prox- imalis and portio distalis are attached to each other by a cartilaginous piece, with this at- taching piece being folded at its outer margin like a cover of a folder so that one part of it becomes dorsal and the other ventral. The dorsal part is described as narrower and is attached to the flattened, curved ante- rior end of the portio proximalis like the blades of scis- sors on its counterpart. The nature of attachment of the two pieces of the anterior cornua renders them capable of opening out to some extent like the covers of a folder. The description of V. monitor (Sondhi 1958) and our dissection of V. indicus (BYU Fig. 13. Hyoid apparatus of Varanus indicus (BYU 40944). Ventral view with the left epihyal reflected to show the absence of a cartilage connection between it and the distal end of the hyoid cornu. Dotted lines ex- tending from the cartilaginous median part of the epi- hyal represents connective tissue. The elongate first epi- b ranch ials are cut. 40944) differ somewhat. We did not find a cartilaginous connection between the portio proximals (hyoid cornu) and the portio dis- talis (epihyal). The only attachment is a later- al sheet of connective tissue that provides a loose connection. The expanded ends are not connected medially and are, therefore, folded as two separate sheets. Near the middle of the epihyal of V. indicus, a thin lateral ex- pansion of cartilage is connected by a sheet of connective tissue to the lateral edge of the hyoid cornu. The distal end of the hyoid cornu is slightly flattened, but not expanded (Fig. 13). The lingual process is shorter than that of V. monitor as figured by Sondhi, and does not extend anterior to the level of the expanded anterior ends of the hyoid cornu and the epi- hyal. In Varanus the first ceratobranchial and first epibranchial are greatly elongated, and the latter taper to a small rod terminating in connective tissue anterodorsal to the shoulder. In Heloderma the second ceratobranchials are lost, and the epihyal is continuous with the hyoid cornu, forming a sigmoid curve. A joint exists at their point of articulation. The first ceratobranchials are also curved and di- verge far laterally at their distal ends. In Xenosaurus as well, the second ceratobran- chials are lost, but the epihyals are straight and long, with a hook at their distal end. The area of articulation between the epihyal and hyoid cornu is enlarged to form a knob. The hyoid cornua extend anterolaterally about two-thirds the length of the lingual process. McDowell and Bogert (1954) report that the hyoids of Lanthanotus and Heloderma are basically similar except LMnthanotus has lost the epihyals. Rieppel (1981) investigated Lanthanotus and found hypohyals (epihyal of McDowell and Bogert) that were reported absent by McDowell and Bogert (1954), al- though McDowell (1972:213) later did report them to be present. Rieppel rejects the argu- ment of McDowell and Bogert that Lantha- notus is close to the origin of snakes. Rieppel (1981:435) states, neither the shape of the basihyal nor any other feature of the hyobranchial skeleton of Lanthanotus shows a particular similarity to the ophidian hyoid. Through the courtesy of Dr. Richard Zwei- fel we were privileged to examine the throat September 1982 Tanner, Avery: Buccal Floor of Reptiles 303 Fig. 14. Hyoid apparatus, ventral views: A, Lanthanotus borneensis (AMNH 87375); B, Heloderma suspectum (BYU 41436). anatomy of iMnthanotus boreensis (AMNH 87375) and found the hyoid skeleton to be smprisingly similar to that of Heloderma (Fig. 14). Rieppel (1980, 1981) has, on the basis of cranial anatomy, concluded that Lanthanotus is intermediate in structure between Helo- derma and Varanus. Branch (1982) arrived at a similar conclusion based on hemipeneal data. The hyoid of these genera have the same structures; however, in Varanus there has been considerable modification and spe- cialization not found in the other genera. In Gerrhosaurus (Cordylidae) the second ceratobranchials have been lost, but the first ceratobranchial and epihyal are retained. In Zonurus the second ceratobranchials are present but short. In Xantusia (Xantusidae) the hyoid contains all the elements. The hyoid comu extends dorsolaterally to articu- late with the median edge of the expanded, flattened proximal end of the epihyal. From the flattened end the epihyal extends post- erodorsally, tapering into a rod and termi- nating as a short epibranchial immediately posterior to the tympanum. The first cerato- branchial extends posterodorsally and curves to terminate in the second epibranchial and in close association with the epibranchial of the epihyal. The second ceratobranchial in Xantusia extends posterior with the distal end, curving laterad to form an open hook. It does not ar- ticulate with an epibranchial as in the epi- hyal and first ceratobranchial; however, a cartilaginous structure in close association with the distal end of the second ceratobran- chial extends laterally and curves anteriorly to articulate with the basioccipital of the skull. Cope (1900) and Savage (1963) have re- ferred to this structure as a free epibranchial. If this is an epibranchial, it is distinct and differs from all others in saurians we have seen. Its close association to the distal end of the second ceratobranchial (Fig. 15) is not ar- ticulated as in the other epibranchials and leads us to believe that the entire structure may represent fusions of other remnant gill bars. An examination of the entire structure (Fig. 15B) indicates to us that fusions have occiirred. An articulation or close association of the distal ends of the epihyal and /or the second ceratobranchial occurs in many forms 304 Great Basin Naturalist Vol. 42, No. 3 Fig. 15. Hyoid apparatus of Xantusia vigilis (BYU 21765): A, ventral view; B, lateral view. (FE = "free epibranchial") but the "free epibranchial" is unique to the xantusids. The hyoid of Anniella has been described by Cope (1892) and Rieppel (1981). Accord- ing to Rieppel, the basihyal is bifurcated pos- teriorly and bears a long entoglossal process. It articulates posteriorly with first cerato- branchials, and small hyohyals (hyoid cornua) are present. These latter structures were con- sidered absent by Cope and Langebartel (1968). In Amphisbaenia all the elements are pres- ent, with the second ceratobranchials being short and widely separated. The hyoid cornu extends anterolaterally, with its distal end free. The epihyal articulates with the cornu about one quarter of its distance from the proximal end. The first ceratobranchial artic- ulates at the point of articulation between the hyoid cornu and the body of the hyoid. Its terminal end bears an epibranchial. All the posterior projections of the hyoid extend straight back and remain unattached at their distal ends (Fig. 16)., Ophidia The hyoids of snakes have been extensively discussed by Langebartel (1968) and others as follows: Anomalepididae Anomalepis (Smith and Warner 1948), Helminthophis (List 1966, Langebartel 1968), Liotyphlops (List 1966, Langebartel 1968). Typhlopidae Typhlophis (Evans 1955, List 1966), Ty- phlops (List 1966, Langebartel 1968). Leptotyphlopidae Leptotyphlops (Smith and Warner 1948, List 1966, Langebartel 1968, Oldham, Smith, and Miller 1970). September 1982 Tanner, Avery: Buccal Floor of Reptiles 305 CBll Fig. 16. Hyoid apparatus, ventral views: A, Amphisbaenia cornura (BYU 16127); B, Amphisbaenia kingi (BYU 16148). Uropeltidae Platyplactrurus (Langebartel 1968), Plec- trurus (Rieppel 1981), Rhinophis (Smith and Warner 1948, Langebartel 1968), Sily- bura (Langebartel 1968). Aniliidae Anilius (Smith and Warner 1948, Lange- bartel 1968, Rieppel 1981), Cylindrophis (Smith and Warner 1948, Langebartel 1968). Xenopeltidae Xenopeltis (Smith and Warner 1948, Langebartel 1968). Boidae Aspidites (Smith and Warner 1948, Lange- bartel 1968), Boa (Langebartel 1968), Calabaria (Langebartel 1968), Charina (Langebartel 1968), Chondropython (Langebartel 1968), Constrictor (Langebar- tel 1968), Enygrus (Langebartel 1968), Epicrates (Langebartel 1968), Liasis (Langebartel 1968), Lichanura (Langebar- tel 1968), Loxocemus (Smith and Warner 1948, Langebartel 1968), Nardoana (Langebartel 1968), Python (Furbringer 1922, Langebartel 1968, Oldham, Smith, and Miller 1970), Sanzinia (Langebartel 1968), Trachyboa (Langebartel 1968). Colubridae Achalinus (Langebartel 1968), Achro- chordus (Smith and Warner 1948, Lange- bartel 1968), Adelphicus (Langebartel 1968), Amblycephalus (Smith and Warner 1948, Langebartel 1968), Aparallactus (Langebartel 1968), Apostolepis (Lange- bartel 1968), Atretium (Langebartel 1968), Boiga (Langebartel 1968), Carphophis (Smith and Warner 1948, Langebartel 1968), Cerberus (Langebartel 1968), Chersodromus (Langebartel 1968), Cher- sydrus (Langebartel 1968), Chrysopelea 306 Great Basin Naturalist Vol. 42, No. 3 (Langebartel 1968), Clelia (Langebartel 1968), Coluber (Walter 1887, Langebartel 1968), Coniophanes (Langebartel 1968), Conophis (Langebartel 1968), Conopsis (Langebartel 1968), Crotaphopeltis (Lang- ebartel 1968), Cyclagras (Langebartel 1968), Dasypeltis (Smith and Warner 1948, Langebartel 1968), Dendrophidion (Langebartel 1968), Diadophis (Langebar- tel 1968), Dipsadoboa (Langebartel 1968), Dispholidus (Langebartel 1968), Dro- mophis (Langebartel 1968), Dry7narchon (Langebartel 1968), Drymobius (Langebar- tel 1968), Dryophis (Langebartel 1968), Elaphe (Langebartel 1968), Elapomorphus (Langebartel 1968), Elapops (Langebartel 1968), Enhydrus (Langebartel 1968), Enulius (Langebartel 1968), Farancia (Langebartel 1968), Ficimia (Langebartel 1968), Fimbrios (Langebartel 1968), Geophis (Langebartel 1968), Haldea (Langebartel 1968), Haplopeltura (Lang- ebartel 1968), Heterodon (Weaver, 1965, Langebartel 1968), Homalopsis (Langebar- tel 1968), Lampropeltis (Langebartel 1968), Leptodeira (Langebartel 1968), Lep- tophis (Langebartel 1968), Manolepis (Langebartel 1968), Masticophis (Lang- ebartel 1968), Mehelya (Langebartel 1968), Natrix (Sondhi 1958), Nerodia (Langebartel 1968, Oldham, Smith, and Miller 1970), Ninia (Langebartel 1968), Nothopsis (Langebartel 1968), Opheodrys (Langebartel 1968, Cundall 1974), Oxy- belis (Langebartel 1968), Oxyrhabdinium (Langebartel 1968), Fituophis (Smith and Warner 1948, Bullock and Tanner 1966, Langebartel 1968, Oldham, Smith, and Miller 1970), Psamaodynastes (Langebar- tel 1968), Rhadineae (Langebartel 1968), Rhadinella (Langebartel 1968), Rhino- cheilus (Langebartel 1968), Salvadora (Langebartel 1968), Sibynomorphus (Langebartel 1968), Sibynophis (Langebar- tel 1968), Sonora (Langebartel 1968), Tan- tilla (Langebartel 1968), Thamnophis (Bullock and Tanner 1966, Langebartel 1968, Oldham, Smith, and Miller 1970), Toluca (Langebartel 1968), Trimorphodon (Langebartel 1968), Tropidonotus (Lang- ebartel 1968), Xenodermus (Langebartel 1968), Xenodon (Weaver 1965). Elapidae Acanthophis (Langebartel 1968), Aspide- laps (Langebartel 1968), Rungarus (Lange- bartel 1968), Calliophis (Langebartel 1968), Demansia (Langebartel 1968), Den- draspis (Langebartel 1968), Denisonia (Langebartel 1968), Doliophis (Langebar- tel 1968), Flaps (Langebartel 1968), Elap- soidea (Langebartel 1968), Furina (Lange- bartel 1968), Hemachatus (Langebartel 1968), Hemibungarus (Langebartel 1968), Leptomicrurus (Langebartel 1968), Mati- cora (Langebartel 1968), Micruroides (Langebartel 1968), Micrurus (Smith and Warner 1968, Langebartel 1968), Naja (Langebartel 1968, Kamal, Hamouda, and Mokhtar 1970), Notechis (Langebartel 1968), Ogmodon (Langebartel 1968), Pseiidelaps (Langebartel 1968), Ultoca- lamus (Langebartel 1968). Eydrophidae Aipysurus (Langebartel 1968), Hydrophis (Langebartel 1968), Kerilia (Langebartel 1968), Lapemis (Smith and Warner 1948, Langebartel 1968), Laticauda (Langebar- tel 1968), Thalasophina (Langebartel 1968). Viperidae Aspis (Langebartel 1968), Atheris (Lange- bartel 1968), Atractaspis (Langebartel 1968), Ritis (Langebartel 1968), Causus (Langebartel 1968), Cerastes (Langebartel 1968), Echis (Langebartel 1968), Pseudoce- rastes (Langebartel 1968), Vipera (Lange- bartel 1968, Furbringer 1922). Crotalidae Agkistrodon (Smith and Warner 1948, Langebartel 1968), Rothrops (Langebartel 1968), Crotalus (Langebartel 1968, Old- ham, Smith, and Miller 1970), Lachesis (Langebartel 1968), Sistrurus (Langebartel 1968), Trimeresurus (Langebartel 1968). In snakes the hyoid apparatus is greatly re- duced, with the hyoid cornua being lost and the remainder of the processes simplified. Es- sentially the snake hyoid consists of a body plus a lingual process and what is thought to be the second ceratobranchials, which are fused to the body of the hyoid (Figs. 17A and B, 29). The variations found in ophidian hyoids have been discussed by Furbringer September 1982 Tanner, Avery: Buccal Floor of Reptiles (1922), Versluys (1936), Gnanamuthu (1937), Smith and Warner (1948), Sondhi (1958), Al- bright and Nelson (1959), List (1966), Under- wood (1967), Langebartel (1968), Rieppel (1981), and others. There are four major mor- phological types that can be distinguished in snakes. Tliese correspond in shape roughly to the letters M, Y, and V, and to a parallel type 11. The most complete survey of the hyoids of snakes is presented by Langebartel (1968), and we have based much of our remarks on his study. Hyoids possessing the M shape are found exclusively in the family Anomalepididae, which has only four genera, Anotnalepis, Lio- typlilops, Hehninthophis, and Tijpldopfiis. In this group tlie hyoid has a body and the sec- ond ceratobranchials. All other processes are lost, including the lingual process. A Y-shaped hyoid is foimd in the Typl- opidae and Leptotyphiopidae. The body of the hyoid possesses a lingual process and has hyoid cornua (second ceratobranchials) that project posteriorly. The possession of a ling- ual process is variable, with it being absent according to List (1966) in TypJiIops pusillus and T. hanbricalis. In T. reticulatiis, T. pla- tycephahis, and T. blandfordi lestradei the hyoid cornua are separated from the body. Leptoti/pJdops has a normal Y type hyoid. Tlie V-shaped hyoid is found in the Ani- liidae, Boidae, Uropeltidae, and Zenopel- tidae. In this type of hyoid the lingual pro- cess is absent and the hyoid cornua may be attached or imattached. There is much in- traspecific variation in the latter character. In some specimens of Charina hottae the cornua are attached, although they are unat- tached in others. Langebartel (1968) consid- ers the curving arches to be the first ceratobranchials. The 11 type hyoid is fomid in the colu- brids, crotalids, elapids, hydrophids, viperids, and some genera of the boidae {Casarea, Tra- cJnjboa, and TropiJopJiis). The second cerato- branchials of this type are usually long, paral- lel rods attached to a slim hyoid body (Fig. 17). The resulting structure resembles a tim- ing fork in appearance. A few snakes have a hyoid body, triradiate in appearance and with a short lingual process. Such a structure is figured by Sondhi (1958) for Natrix (Xe- nochrophis), in which: 307 Bh CBI 4X 1^ B 4X Fig. 17. Hyoid apparatus, ventral views: A, Pituophis m. deserticola (BYU 3072); B, Crotahts viridis lutosus (2089). Both are from adult individuals and drawn at 4X actual size. 308 Great Basin Naturalist Vol. 42, No. 3 tilt' l)asihvoid lies ventral to the trachea and dorsal to the posterior terminations of the oniohyoideus and ster- nohvoidens muscles. The processes form elongated rods that lie ventral and extend posteriorly and parallel, with their terminal ends enclosed in the tips of the base of the tongue. In Pitiiophis, the basihyoid is ventral to the tongue at about the level of the angle of the jaws. The cerato- branchials extend and curve posterolaterally from the basihyoid for a short distance to a lateral position and then extend posteriorly, lateral to the tongue and parallel to each other, to the posterior tip of the tongue. In Crotalus the same obtains anteriorly with the basihyoid and the anterior part of the cerato- branchials; however, the posterior third of the latter converge ventrally to become closely associated along the ventromedian of the tongue and diverge slightly near their ends to become imbedded in muscle and con- nective tissue (Fig. 28, Romer 1950: fig. 421- C, Langebartel 1968:figs. 3, 4). For detailed description of the hyoid of individual genera of snakes, see Langebartel (1968). III. Muscles of the Buccal Floor: General The buccal floor is composed of several in- terwoven sheets of muscles. These sheets can be separated into two major groups: the hypobranchial musculature and the muscles of the associated branchial arches. The hypo- branchial muscles are derived from the myo- tomes of the occipital and cervical somites, whereas the muscles of the branchial arches come from tlie viceral muscle plates formed in tlie branchial region. The tongue, for the most part, is also derived from the occipital somites. Because of the close associations of some of the somites with both cranial and spinal areas, some muscles are innervated by both spinal and cranial nerves. For the sake of convenience, we have sepa- rated our discussion of the buccal muscula- ture into two major divisions: (1) the muscles associated with the hyoid apparatus and (2) those associated with other structures. The tongue is svifficiently important to be segre- gated from these categories and is considered under a separate heading. The nomenclature of muscles of reptiles has not been standardized; however, tables of synonyms can be found in Edgeworth (1935), Langebartel (1968), Haas (1973), and Schumacher (1973). Some of the more recent short summaries of the earlier papers on the myology of the buccal floor in reptiles can be found in Sondhi (1958), Langebartel (1968), Avery and Tanner (1971), Secoy (1971), and Varkey (1979). The remainder of this section is a brief account of the musculature of the buccal floor in selected reptiles as described by several earlier workers such as Edgeworth (1931), Graper (1932), Gnanamuthu (1937), Reese (1915 and 1932), Hacker and Schuma cher (1955), Oelrich (1956), Sondhi (1958), Langebartel (1968), and others. It also should be noted that the more advanced reptiles have more complex muscular patterns when compared to primitive forms. This is seem- ingly true not only for orders, but also for family groups. A comparison of the advanced lizard Varanus and the primitive iguanids in the following sections serves as an illustration. We refer to such forms as Gavialis, Tri- onyx, Natrix (Xenochropliis), Varanus, and other genera. These should be credited to Gnanamuthu (1937) or Sondlii (1958) if not otherwise noted. The musculature of the following reptiles has been studied. Chelonia Pelomedusidae Pelusios (Poglayen-Neuwall 1953a). Chelidae Batrochemys (Poglayen-Neuwall 1953a), Chelodina (Poglayen-Neuwall 1953a). Chelydridae Chelydra (Camp 1923, Graper 1932, Pog- layen-Neuwall 1953a, Schumacher (1973), Kinosternon (Poglayen-Neuwall 1953a, Schumacher 1973), Sternotherus (Pog- layen-Neuwall 1953a, Schumacher 1973). Testudinidae Chrysemys (Poglayen-Neuwall 1953a, Ash- ley 1955, Schumacher 1973), Cuora (Pog- layen-Neuwall 1953a), Clemmys (Graper 1932), Poglayen-Neuwall 1953a, Schuma- cher 1973), Deirochelys (Shah 1963), Emys September 1982 Tanner, Avery: Buccal Floor of Reptiles 309 (Walter 1887, Schumacher 1973), Goph- erus (George and Shad 1954), Graptemys (Poglayen-Neuwall 1953a), Geochelone (Bojanus 1819, Graper 1932, Lubosch 1933, Edgeworth 1935, Poglayen-Neuwall 1953a, George and Shad 1955, Schuma cher 1973), Malaclemys (Poglayen-Neu- wall 1953a), Pseudemys (Ashley 1955, Poglayen-Neuwall 1953a, Schumacher 1973), Terrapene (Poglayen-Neuwall 1953a). Trionychidae Lissevnjs (George and Shad 1954, Sondhi 1958, Schimiacher 1973), Trionyx (Graper 1932, Lubosch 1933, Poglayen-Neuwall 1953a, Sondhi 1958, Schumacher 1973). Cheloniidae Caretta (Poglayen-Neuwall 1953a, Schumacher 1973). Dermochelyidae Dennochelys (Poglayen-Neuwall 1953a, 1953/54, Schumacher 1973). Rhynchocephalia Sphenodontidae Sphenodon (Osawa 1898, Camp 1923, Byerly 1926, Lubosch 1933, Edgeworth 1935, Lightoller 1939, Kesteven 1944, Rieppel 1978). Lacertilia Gekkonidae Coleonyx (Camp 1923), Gekko (Camp 1923, Lubosch 1933), Gymnodactylus (Brock 1938, Kesteven 1944), Hemidac- tylus (Zavattari 1908, Ping 1932, Edge- worth 1935, Gnanamuthu 1931), Platydac- tylus (Sanders 1870, Poglayen-Neuwall 1954), Stenodactylus (Camp 1923, Edge- worth 1935), Tarentola (Gnanamuthu 1937, Poglayen-Neuwall 1954), Thecodac- tyhis (Kesteven 1944). Dibamidae Dihamua (Case 1968). Iguanidae AmbJrhynchus (Avery and Tanner 1971), Anolis (Kesteven 1944), Basiliscus (Gnana- muthu 1937), Brachylophus (Gamp 1923, Avery and Tanner 1971), Callisaiirus (Cox and Tanner 1977), Chalarodon (Avery and Tanner 1971), Conolophus (Cox and Tan- ner 1977), Crotaphytus (Davis 1934, Rob- ison and Tanner 1968), Ctenosaura (Oel- rich 1956, Avery and Tanner 1971), Cyclura (Avery and Tanner 1971), Dipso- souriis (Avery and Tanner 1971), Enyalio- saurus (Avery and Tanner 1971), Hoi brookia (Cox and Tanner 1977), Iguana (Mivart 1867, Edgeworth 1935, Poglayen- Neuwall 1954, Avery and Tanner 1971, Oldham and Smith 1975), Oplurus (Avery and Tanner 1971), Phrynosoma (Sanders 1874, Camp 1923, Jenkins and Tanner 1968), Sauromahis (Avery and Tanner 1964, 1971), Scehporus ^Secoy 1971), Tropidurus (Zavattari 1908, Edgeworth 1935), Uma (Cox and Tanner 1977), Uro- saiirus (Fanghella, Avery and Tanner 1975), Uta (Fanghella, Avery and Tanner 1975). Agamidae Agama (DeVis 1883, Lubosch 1933, Edge- worth 1935, Poglayen-Neuwall 1954, Har- ris 1963), Amphibolurus (Poglayen-Neu- wall 1954), Calotes (Camp 1923, Gnanamuthu 1937, Poglayen-Neuwall 1954), Chlamydosaurus (DeVis 1883), Draco (Gnanamuthu 1937), Leiolepis (Sanders 1872, Poglayen-Neuwall 1954), Phrynocephalus (Kesteven 1944), Phys- ignathus (Kesteven 1944), Sitana (Gnana- muthu 1937), Uromastix (Furbringer 1922, Kubosch 1933, Edgeworth 1935, George 1948, Poglayen-Neuwall 1954, Throck- morton 1978). Chamaeleonidae Chamaeleo (Mivart 1870, Zavattari 1908, Camp 1923, Lubosch 1933, Edgeworth 1935, Gnanamuthu 1937, Kesteven 1944, Poglayen-Neuwall 1954). Scincidae Eumeces (Zavattari 1908, Edgeworth 1935, Nash and Tanner 1970), Mabuya (Gnanamuthu 1937), Tiliqua (Lightoller 1934, Kesteven 1944, Poglayen-Neuwall 1954), Trachysaurus (Poglayen-Neuwall 1954). Cordylidae Cordylus (Camp 1923, Edgeworth 1935), Gerrhosaurus (Camp 1923). 310 Great Basin Naturalist Vol. 42, No. 3 Lacertidae Cabrita (Gnanamuthu 1937), Lacerta (Walter 1887, Camp 1923, Edgeworth 1935, Poglayen-Neuwall 1954). Teiidae Ameiva (Poglayen-Neuwall 1954, Fisher and Tanner 1970), Cnemidophorus (Pog- layen-Neuwall 1954, Fisher and Tanner 1970), Tupinambis (Zavattari 1908, Camp 1923, Edgeworth 1935, Poglayen-Neuwall 1954). Anguinidae GerrJwnotus (Camp 1923, Poglayen-Neu- wall 1954), Ophiosaurus (Poglayen-Neu- wall 1954). Xenosauridae Shinosaurus (Haas 1960), (Camp 1923, Haas 1960). Xenosaurus Helodermatidae Heloderma (Camp 1923, Poglayen-Neu- wall 1954). Varanidae Varanus (Bradley 1903, Camp 1923, Edgeworth 1935, Gnanamuthu 1937, Lightoller 1939, Kesteven 1944, Poglayen- Neuwall 1954, Sondhi 1958). Anniellidae Anniella (Camp 1923). Amphisbaenidae Amphisbaena (Smalian 1885, Camp 1923), Anopsibaena (Smalian 1885). Bipes (Sma- lian 1885, Renous 1977), Blanus (Smalian 1885), Rhineura (Camp 1923), Trogo- nophis (Smalian 1885). Xantusidae Xantusia (Camp 1923). Ophidia Anomalopididae Anomalepis (Haas 1968), Hehninthophis (Langebartel 1968), Liotyphlops (Lange- bartel 1968). Typhlopidae Typhlophis (Evans 1955), Typhlops (Langebartel 1968). Leptotyphlopidae Leptotyphlops (Langebartel 1968, Oldham, Smith, and Miller 1970). Uropeltidae Plutylectrurus (Langebartel 1968), Rhi- nophis (Langebartel 1968), Uropeltis (Langebartel 1968). Aniliidae Anilius (Langebartel 1968), Cylindrophis (Lubosch 1933, Langebartel 1968). Xenopeltidae Xenopeltis (Langebartel 1968). Boidae Boa (Gibson 1966), Calabaria (Langebartel 1968), Charina (Langebartel 1968), Con- strictor (Langebartel 1968), Epicrates (Langebartel 1968), Eryx (Langebartel 1968), Eunictes (Anthony and Serra 1950, Langebartel 1968), Liasis (Langebartel 1968), Python (Lubosch 1933, Edgeworth 1935, Kesteven 1944, Frazzetta 1966, Langebartel 1968, Oldham, Smith, and Miller 1970), Sanzinia (Langebartel 1968), Trachyboa (Langebartel 1968). Colubridae Achalinus (Langebartel 1968), Achro- chordus (Langebartel 1968), Amblyce- phalus (Langebartel 1968), Aparallactus (Langebartel 1968), Atretium (Langebartel 1968), Cerfoems. (Langebartel 1968), Cher- sydrus (Langebartel 1968), Coluber (Wal- ter 1887), Dasypeltis (Langebartel 1968), Dryophis (Lubosch 1933), Elaphe (Albright and Nelson 1959, Langebartel 1968), En- hydrus (Langebartel 1968), Fimbrios (Langebartel 1968), Haplopeltura (Lange- bartel 1968), Heterodon (Langebartel 1968), Mehylya (Langebartel 1968), Matrix (Sondhi 1958), Nerodia (Langebartel 1968, Oldham, Smith, and Miller 1970, Varkey 1979), Nothopsis (Langebartel 1968), Oph- eodrys (Cundall 1974), Pituophis (Oldham, Smith, and Miller 1970), Sibynomorphus (Langebartel 1968), Sibynophis (Langebar- tel 1968), Thamnophis (Langebartel 1968, Oldham, Smith, and Miller 1970), Tropido- notus (Lubosch 1933), Xenodermus (Langebartel 1968), Xenodon (Langebartel 1968). September 1982 Tanner, Avery: Buccal Floor of Reptiles 311 Elapidae Denisonia (Langebartel 1968), Doliophis (Langebartel 1968), Naja (Lubosch 1933, Langebartel 1968), Notechis (Langebartel 1968), Pseudechis (Kesteven 1944). Hydropidae Aipysurus (Langebartel 1968), Hydrophis (Langebartel 1968), Laticauda (Langebar- tel 1968), Pelamis (Langebartel 1968). Viperidae Aspis (Langebartel 1968), Atractaspis (Langebartel 1968), Caiisus (Haas 1952, Langebartel 1968), Cerastes (Langebartel 1968), Echis (Langebartel 1968), Vipera (Edgeworth 1935, Langebartel 1968). Crotalidae Agkistrodon (Langebartel 1968, Kardong 1973), Bothrops (Langebartel 1968), Cro- talus (Langebartel 1968, Oldham, Smith, and Miller 1970), Lachesis (Lubosch 1933, Langebartel 1968). Crocodilia Crocodylidae Alligator (Reese 1915, Lubosch 1933, Edgeworth 1935, Chiasson 1962, Pog- layen-Neuwall 1953b), Caiman (Schuma- cher 1973), Crocodylus (Camp 1923, Edge- worth 1935, Kesteven 1944, Sondlii 1958, Poglayen-Neuwall 1953b). Gavialidae Gavialis (Sondhi 1958). IV. Buccal Floor Muscles Associated WITH THE HyOID APPARATUS 1. M. geniohyoideum (genioglossus) The M. geniohyoideus originates on the mandible and inserts on the hyoid apparatus. In Lissemys the M. geniohyoideus consists of two bimdles arising from the mandible and inserting on the second ceratobranchial. Two distinct parts of this muscle arise from sepa- rate although continuous sites on the man- dible in Trionyx. Each part inserts individ- ually on the second ceratobranchial. According to Sondhi (1958) one of these, the portio dorsalis, arises from the ventral surface of the second ceratobranchial. The other, the portio ventralis, lies ventral to the portio dor- salis and dorsal to the Mm. mylohyoideus posterior and constrictor colli; it inserts on the second ceratobranchial just posterior to the portio dorsalis. In Deirochelys and Chelodina one part (M. genioglossus) arises from the anterior end of the inner border of the dentary and inserts on the basihyoid. Another portion (M. gen- iohyoideus) arises from the inner side of the mandibular symphysis and passes posteriorly to insert on the proximal end of the hyoid comua. A similar condition exists in Lissemys and Geochelone elegans except that the me- dian fibers also insert on the median raphe. The M. geniohyoideus oi Alligator is a slen- der muscle separated into two bundles. The medial bundle inserts onto the second cerato- branchial, whereas the lateral attaches to the M. sternohyoideus. The M. geniohyoideus of Gavialis lies obliquely in the posterior part of the buccal floor, where it originates posteri- orly along the inner border of the mandible; it extends posteriorly and medially to become a tendon at its insertion near the middle of the ventrolateral border of the ceratobran- chial. In Crocodylus the M. geniohyoideus in- serts on the ventrolateral aspect of the prox- imal part of the second ceratobranchial. In Sphenodon (Byerly 1926) and Cha- maeleo (Gnanamuthu 1937) it is narrow, whereas in Mahtiia, Cabrita, Anolis (Gnana- muthu 1937), Amblyrhynchus, Brachylophus, Chalarodon, Conolophus, Ctenosaura, Cy- clura, Dipsosaurus, Iguana, Oplurus, Sauro- malus (Avery and Tanner 1971), Hemidac- tylus, Coleonyx, Tarentola (Figs. 4, 5), Chlamydosaurus (Beddard 1906), Uromastyx, Xenosaurus (Haas 1960), Cnemidophorus (Fisher and Tanner 1970, Presch 1971), Helo- derma, Gerrhonotus (Camp 1923), Anniella (Bellairs 1950), Shinisaurus (Haas 1960), and Dibamus (Girgis 1961, Case 1968) it forms a broad sheet arising from the posteromedial border of the mandible and passing posteri- orly. There it is divided into three to six slips that may interdigitate with the M. mylo- hoideus (Fig. 18 A, B, C, D). The superficial lateral slips overlie the medial one posteri- orly and insert on the first ceratobranchial ventral to the medial muscle. A deep lateral slip originates on the mandible dorsal to the 312 Great Basin Naturalist Vol. 42, No. 3 Fig. 18. Ventral view of the superficial supporting muscles of the throat and buccal floor: A, the gecko Tarentola annularis (BYU 18122); B, Sceloponis magister (BYU 30310); C, Ameiva n. parva (BYU 14396); and D, Tarentola with superficial muscles removed. The closely adhering skin in Sceloponis shows the scale impressions. Gh-Geniohyoideus; Mha-Mylohyoideus anterior; Prh-Prearticulohvoideus; Mhp-Mylohyoideus posterior; Ptm-Pterygomandibularis; Oh- Omohyoideus. i September 1982 Tanner, Avery: Buccal Floor of FIeptiles 313 Fig. 19. Ventral view of Phrynosoma platijrhinos: A, Superficial myology; B, deeper muscles. (After Jenkins and Tanner 1968) lateral superficial slip and inserts on the dis- tal end of the epihyal. The geniohyoideus in Varaniis arises from the ventromedial border of the posterior part of the mandibular ramus and fans out posteri- orly to cover the buccal floor and neck. The fibers converge posteriorly to insert on the ventromedial border of the proximal end of the second ceratobranchial and basihyoid, and the more median fibers insert in the fas- cia of the stemohyoideus and omohyoideus muscles. In the Iguanidae the medial fibers insert on the basihyoid or the anterior margin of the first ceratobranchials (Fig. 19), where- as in the gekkonids {Tarentola and Coleonyx) fibers are loosely divided into two bundles, the irmer one inserting on the basihyoid and the other attached along the anterior margin of the first ceratobranchial (Fig. 18 D). The M. geniohyoideus (genioglossus of Av- ery and Tanner 1971) of the iguanine lizards consists of three parts, including the anterior fibers that arise on the ventromedial border of the mandible, where its fibers interdigitate with the M. intermandibularis anterior pro- fundus and extend posteriorly (Fig. 18). There, the more medial fibers may insert on the lingual process, with the remainder pass- ing ventral to the anterior comu to insert on the first ceratobranchial (Fig. 19). A second division originates on the midventral raphe and inserts on the anterior border of the first ceratobranchial, with the third portion origi- nating on the ventromedial border of the mandible, and interdigitates (as does the first part) before inserting on the lateral border of the first ceratobranchial. A muscle deep to the lateral slip originates on the mandible and inserts on the posterior edge of the epi- hyal. This muscle may easily be included as a part of the lateral slip of the geniohyoideus. Jenkins and Tanner (1968), following Oelrich (1956), referred to it as the M. mandibulo- hyoideus III (Fig. 20). We have modified their designation to the M. mylohyoideus III, and wonder if the muscle is not a part of the M. geniohyoideus adapted to strengthen the lateral part of the mandibular-hyoid-buccal floor. We note that the same muscle is pres- ent in Agama, but less massive than in iguanids. In the scincid Eumeces (Nash and Tanner 1970), the M. geniohyoideus originates from the anteromedial fifth of the mandible and inserts posteriorly by medial and lateral slips onto the hypoglossus, lingual fascia, and ante- rior margin of the first ceratobranchial. Some fibers also insert dorsally on the oral mem- brane and anteromedially on the cutaneous fascia. Fisher and Tanner (1970) describe the M. geniohyoideus in Ameiva and Cnemidophorus (Teiidae) as originating on the medial surface of the dentary and inserting as five slips along the anterior margin of the body of the hyoid and the first ceratobranchial. Some dorsal fibers appear to insert on the ventral portion of the tongue. In these genera there is considerable interdigitation of the trans- verse and longitudinal muscles, as seen in Figure 18 C. 314 Great Basin Naturalist Vol. 42, No. 3 GHII Fig. 20. Ventral view of the M. geniohyoideus of Sauromalus (BYU 32551) showing the origins (along mandible) and insertions (on hyoid apparatus). Gh-I-II- III divisions of the genioglossus and Mh III mandibulohyoideus. In Calotes, Sitana, and Chamaeleo (Fig. 18) the median bundle is similar to that of the geckos, but there are two lateral bundles in Calotes and Chamaeleo and four in Sitana. In Chamaeleo the two lateral bundles are deep- er and insert on the ceratobranchial. The most medial of these bundles also has an in- sertion on the anterior cornu. In Draco there are four lateral bundles, but the median bundle is missing. One of the lateral bundles interweaves with the M. mylohyoideus ante- rior and another (M. geniohyoideus basibran- chialis of Gnanamuthu 1937) is attached to the branchial process. The lateral bundles of the M. geniohyoideus of Chamaeleo and Draco produce the M. adductor inferior la- bioris of Gnanamuthu (1937) (Fig. 21). In Agama agama the median fibers do not insert on the basihyoid, but extend ventral to it and insert on the first ceratobranchial. The anterior cornu and body of the hyoid are cov- ered ventrally by the M. geniohyoideus. The deep lateral slip inserts on the epihyal and, except for its smaller size, is similar to that seen in the iguanids. In snakes such as the anomalepidids, the M. geniohyoideus arises from the posterior half of the mandible and passes posteriorly as a broad sheet separated medially from its counterpart by the linea alba. It inserts on both the basihyal and the second ceratobran- chial. In the anomalepidids a slender slip of muscle attaches to the tip of the dentary and the terminal part of the second ceratobran- chial; it has been described by Langebartel (1968) as being either another portion of the M. geniohyoideus or the M. ceratomandibu- laris. In the anomalepidids there is some vari- ation in this muscle. The origin is by a single head in the Leptotyphlopidae and in the gen- era Rhinophis, Cylindrophis rtifus, Sanzinia, Enhydris, Aidpysurus, and Bothrops. There is more than one head of origin in the Typhlo- pidae and Uropeltidae. Another portion of this complex (M. gen- iohyoideus of Langebartel 1968) is described as occurring only in the Anomalepididae, in which it originates from the posterior half of the lower jaw and inserts on the hyoid cornua and ceratobranchial. In Matrix (Xenochrophis) the M. geniohyoideus is covered ventrally by the Mm. mylohyoideus posterior and con- strictor colli after arising from the ventrome- dial border of the mandible. The parallel fi- bers of the M. geniohyoideus insert on the lateral border of the ba.sihyoid and the ante- rior border of the second ceratobranchial af- ter passing obliquely to the midline. Varkey (1979) describes a second origin from the midventral raphe and fascia just anterior to September 1982 Tanner, Avery: Buccal Floor of Reptiles 315 Fig. 2L Ventral view of Chamaeleon brecicornis (BYU 12422): A, superficial and muscles immediately dorsal to the superficial ones; B, geniohyoideus. the lingual sheath. He considers the insertion to be the fascia of the hypoglossus muscle. 2. M. genioceratoideus In Varanus the most lateral bundles of the M. geniohyoideus complex form a separate muscle (Sondhi 1958). It arises on the inner ventrolateral border of the mandible and ex- tends posteriorly, where its fibers divide into two bundles. One bundle inserts on the later- al side of the handlelike position of the portio proximalis of the anterior comu, with the second bundle inserting on the ventrolateral border of the middle cartilaginous part of the portio distalis of the second ceratobranchial. This muscle may exist in Chamaeleo, in which it has been described by Mivart (1870) as the ceratomandibular. A similar situation exists in Chhmydosaurus (Beddard 1950b, DeVis 1883). 3. M. prearticulohyoideus The M. prearticulohyoideus is considered as a division of the M. genioceratoideus by Gnanamuthu (1937). 3a. M. mandibulohyoideus In turtles, such as Trionyx, this muscle is large, lying in the ventrolateral region of the buccal floor and arising from the ventrome- dial border of the posterior part of the man- dible; it inserts on the posterior region of the second ceratobranchial. In Gavialis the M. prearticulohyoideus is a thin muscle lying dorsal to the M. ceratohyoideus to insert on the posterior portion of the second cerato- branchial. Edgeworth (1935) has described a similar muscle in Alligator, which he calls the M. branchiomandibularis. The second sheet (M. mandibulohyoideus I) is a long triangular muscle extending two- thirds the length of the mandible and lyin^ lateral to the M. mandibulohyoideus II. This sheet originates along the ventromedial sur- face of the dentary and a small portion of the angular, with some fibers interdigiting with the more superficial musculature. The in- sertion is just posterolateral to that of the M. mandibulohyoideus II on the distal two-thirds of the posterior cornu. 316 Great Basin Naturalist Vol. 42, No. 3 The M. mandibulohyoideus, as described by Avery and Tanner (1971) for the iguanine hzards, by Robison and Tanner (1962) for Crotaphytus, and by Jenkins and Tanner (1968) for Phrynosoma, consists of two sheets. The most medial portion (M. mandibulo- hyoideus II) consists of a pair of small elon- gated bundles of fibers lying medial to the M. mandibulohyoideus I and inserting together on the midventral raphe of the throat. It originates as a narrow tendon from the man- dibular symphysis. Each muscle inserts on the anterior border of the proximal end of the posterior comu. In Varanus Sondhi (1958) described it as a short muscle lying on the ventrolateral side of the neck, covering the posterior part of the mandible ventrally. It arises from the posterior and medial aspects of the mandible and extends almost straight back along the ventrolateral side of the neck to insert on the rodlike portion of the portio distalis of the anterior comu. 4. M. mandibulopriximalis The mandibulopriximalis has been de- scribed in Varanus by Sondhi (1958) as a slender muscle situated dorsal to the M. gen- iohyoideus and ventral to the M. gen- ioglossus. It arises from the ventrolateral bor- der of the ramus of the mandible, extending posteriorly and obliquely to pass dorsal to the handlehke portion of the portio distalis. Most of this muscle inserts on the outer margin of the handlelike portion of the portio prox- imalis, with some of its fibers becoming sepa- rated from the remainder and inserting on the lining of the buccal floor. In the iguanid lizards the M. mandibulo- priximalis, if present, forms a part of the M. geniohyoideus and cannot be distinguished from the latter muscle. 5. M. ceratohyoideus The M. ceratohyoideus of Sphenodon is short and thin, having its origin on the second ceratobranchial and its insertion on tlie ante- rior comu. Rieppel (1978) states that the presence of the M. ceratohyoideus lying be- tween the ceratohyal and the first cerato- branchial and innervated by the M. glos- sopharyngeus is primitive. He further argues that its failure to reach the lower jaw, as is the case in most lizards, is also perhaps an in- dication of its primitiveness. In Lissemys the M. ceratohyoideus arises from the second ceratobranchial and inserts on the basihyoid. In Trionyx it arises from the distal half of the second ceratobranchial, en- closing this cartilaginous rod and extending anteromedially on the lateral side of the buc- cal floor to insert on the middle and anterior components of the basihyoid and on the knoblike anterior comu. In Alligator and Crocodylus it originates on the second ceratobranchial and inserts on the basihyoid. In Gavialis the M. ceratohyoideus lies dorsal to the basihyoid and is not visible in ventral view. The origin is on the dorsola- teral border of the posterior comu, with the muscle extending obliquely forward as a thin sheet on the ventral side of the buccal floor to insert on the dorsolateral side of the ante- rior comu. The insertion in Sitana is on the basihyoid. In Varanus it lies dorsal to the M. gen- iohyoideus on the ventrolateral side of the middle of the neck. The origin is from the ventrolateral border of the proximal piece of the second ceratobranchial, from which the muscle extends anteromedially to fan out over the ventrolateral side of the neck and insert on the handlelike portion of the portio distalis. In the iguanid lizards this muscle has been described as the M. branchiohyoideus by Av- ery and Tanner' (1971). In Ctenosaura the muscle is ribbonlike and situated between the first ceratobranchial and second ceratobran- chial of each side of the hyoid apparatus. The origin is along most of the anterior two-thirds of the first ceratobranchial, with the insertion on the posterior half of the second cerato- branchial. This pattern is duplicated in Cha- larodon. Opiums, Crotaphytus, and all the re- maining iguanine lizards except Sauromalus. In the latter the insertion is very narrow, by a single tendon from the proximal rim of the anterior border of the posterior comu. In Phrynosoma this muscle covers nearly the entire area between the anterior and pos- terior cornua of the hyoid (Fig. 19 B). Its ori- gin and insertion are similar to that described above for other iguanids. In Chamaeleo the M. ceratohyoideus is a small thick mass aris- September 1982 Tanner, Avery: Buccal Floor of Reptiles 317 ing from the posterolateral border of the basihyoid to pass anterodorsally and insert on the epihyal. In Eumeces (Scincidae) the muscle is a nar- row strap similar to that in the iguanid Sauromalus. In the teiids, Cnemidophoms and Ameiva, this muscle has a similar origin to that of the iguanids, but fills the entire area between the anterior and posterior comua (Fisher and Tanner 1970). 6. M. cornuhyoideus The M. cornuhyoideus was described in Varanus by Sondhi (1958) as being immedi- ately posterior to the M. ceratohyoideus; it is ventrally concealed by the basal branch of the tongue and extends between the anterior and posterior comua. It arises from the ven- trolateral border of the proximal piece of the second ceratobranchial and proceeds forward to insert on the outer margin of the portio proximalis of the anterior cornu, anterior to the latter's articulation with the basihyoid. This muscle has not been described in any other reptile. 7. M. interportialis Sondhi (1958) has reported that in Varanus this slender muscle lies dorsal to the M. ce- ratohyoideus and ventral to the portio prox- imalis. The origin is on the ventrolateral side of the anterior portion of the portio prox- imalis, from which the muscle extends obliquely anteriorly to insert on the medial border of the handlelike portion of the portio distalis. Gnanamuthu (1937) did not describe this muscle for Varanus and probably consid- ered it to be part of the M. ceratohyoideus. It has not been described in other reptiles. 8. M. hypoglossolateralis The M. hypoglossolateralis has been de- scribed by Sondhi (1958) as a dehcate strip of muscle lying above the hypoglossum of the tiutle Trionyx. Its origin is on the dorsal sur- face of that cartilaginous plate from which it extends to tlie lining of the buccal floor on which it inserts. This muscle is also present in Gopherus agassizi, and we suspect its pres- ence in association with the hypoglossal car- tilage of other Chelonia. 9. M. entoglossohypoglossalis The M. entoglossohypoglassalis is another muscle described by Sondhi (1958) for Tri- onyx. It arises from the ventrolateral border of the anterior part of the lingual process and inserts dorsolaterally on the posterior surface of the hypoglossum. 10. M. omohyoideus In turtles such as Lissemys the M. omo- hyoideus is thick and long, and has an ante- rior division into dorsal and ventral bundles. The dorsal bundle inserts on the medioprox- imal part of the first ceratobranchial, and the ventral bundle inserts on the basihyoid along with the M. sternohyoideus. In Trionyx the M. omohyoideus originates on the anterior border of the scapula and extends forward on the ventral side of the neck to converge ante- riorly to form two bundles, a larger medial and small lateral, which insert on the pro- ximal part of the second ceratobranchial. In Chelodina the M. omohyoideus arises from the middle of the coracoid, but in Deiro- chelys, Lissemys, and Geochelone it origi- nates on the ventral end of the coracoid. In all genera the fibers pass anteriorly to insert on the ceratobranchials. In Alligator the M. omohyoideus is a long, narrow, thick muscle that originates from the upper border of the coracoid and passes for- ward to insert on the middle of the second ceratobranchial. In Crocodylus the origin is from the anterior border of the scapula and the insertion on the second ceratobranchial. Gavialis, as described by Sondhi (1958), has a moderately broad muscle arising from the an- terior border of the coracoid. As it passes an- teriorly, it divides into two parts, a portio dorsalis and a portio ventralis. The portio dorsalis extends obliquely anteromedially as a narrow strap that terminates in fragile slips that merge into the tendon of the M. ster- nohyoideus. The portio ventralis is broad, and its fibers parallel the trachea, finally in- serting on the short anterior part of the sec- ond ceratobranchial. 318 Great Basin Naturalist Vol. 42, No. 3 The M. omohyoideus is a large muscle that usually arises on the pectoral girdle and in- serts on the hyoid. In Sphenodon it is a large sheet, but in Varanus it is slender and partly covered by the M. stemohyoideus along its medial border. We summarize from Gnana- muthu (1937:24) as follows: In Varanus it arises on the anterior border of the scapula to pass obliquely forward and insert on the an- terior part of the proximal end of the second ceratobranchial close to its articulation with the basihyoid. A similar situation exists in Hemidactylus. In Cabrita, Mahuia, and Cha- maeleo the insertion of the M. omohyoideus is on the anterior border of the basihyoid. In Calotes it inserts not only on the basihyoid, but also on the sides of the proximal part of the first ceratobranchial. In Anolis and Si- tana it inserts only on the first ceratobran- chial, but in Draco there are three bundles, two of which insert on the first ceratobran- chial and the third on the second ceratobran- chial. In Chlamydorsaurus it originates on the clavicle and sternum and inserts on the posterior one-third of the ceratobranchial. In the iguanid lizards, such as Ctenosaura, the M. omohyoideus has medial and lateral origins. Medially the fibers originate on the lateral tip of the transverse process of the in- terclavicle, whereas the lateral fibers origi- nate on the lateral half of the anterolateral surface of the clavicle and the anterior bor- der of the suprascapula. As the two bundles extend anteriorly they become continuous and insert together along the posterior edge of the second ceratobranchial. In all the iguanine lizards and Opiums the fibers of the medial and lateral bundles are impossible to separate. Chalarodon shows a slightly differ- ent configuration, with both bundles being separated for their entire length. The M. omohyoideus in the teiids Ameiva and Cnemidophorus is a thick muscle originating on the anterior border of the sca- pula and then proceeding anteroventrally to insert on the proximal end of the basihyoid and along the second ceratobranchial. In Di- bamus it is extremely long, originating on the scapula and inserting on the distal two-thirds of the ceratobranchial. In snakes this muscle is very small and passes anteriorly from its origin on the lateral body muscles just posterior to the distal end of the hyoid apparatus to insert on the poste- rior portion of the ceratobranchials. It has been found in the Anomalepididae Cylin- drophis, Rhinophis, and Eryx c. colubrinus. 11. M. stemohyoideus The M. stemohyoideus is a complex of muscles that arises from the sternum and in- serts on the hyoid in most reptiles (Fig. 19). In both Lissemys and Trionyx the M. ster- nohyoideus is large and lies adjacent to the M. omohyoideus. It originates from the cla- vicle and passes anteriorly and medially to insert on the proximal part of the second ce- ratobranchial and the middle of the basi- hyoid. In Crocodylus the M. stemohyoideus has long tendons by which it inserts on the second ceratobranchials. In Alligator it is flat and broad, and originates from the ventral surface of the episternum and forms a short tendon that inserts on the M. geniohyoideus. In Gavialis the M. stemohyoideus is a broad flat muscle with an origin on the ventral an- terior half of the episternum; it passes along the ventral side of the neck to meet its oppo- site member at the midline where it obscures the trachea ventrally (Sondhi 1958). As it ap- proaches the hyoid apparatus it divides into two parts, with the outer part (portio ex- terna) a broad band forming a large tendon that inserts on the inner border of the man- dible. The inner bundle (portio interna) par- allels the trachea to insert on the outer part of the posterior border of the basihyoid. In Sphenodon it is flat, whereas in some lizards it becomes cordlike and inserts (Riep- pel 1978) on the caudodorsal edge and dorsal surface of the first ceratobranchial, deep to and lateral to the insertion of the omo- hyoideus. In Mabuia, the M. stemohyoideus inserts on the basihyoid, whereas in Anolis, with its small basihyoid, the insertion is on the first ceratobranchial. In Varanus the M. stemohyoideus lies dorsal to the M. con- strictor colli and ventral to the basihyoid and the proximal piece of the second ceratobran- chial. It arises from the ventrolateral border of the clavicle and extends obliquely ante- riorly to the ventral side of the neck, where it parallels the M. omohyoideus and inserts on the ventral side of the basihyoid and posteri- or portion of the lingual process. Chamaeleo September 1982 Tanner, Avery: Buccal Floor of Reptiles 319 has a small lateral bundle of fibers that insert on the fascia of the lateral M. geniohyoideus. The M. stemohyoideus of the iguanine liz- ards (Avery and Tanner 1971), is an extensive muscle sheet occupying a large area posterior to the anterior comu and anterior to the ster- num and clavicle. It originates from several heads on the clavicle, and its oblique fibers extend anteriorly to insert on the posterior surface of the anterior cornu. In all the iguanines and in Cfialarodon the muscle ap- pears broad and sheetlike. In Opiums, it is narrow and cordlike. In Phrynosorna the M. stemohyoideus is separated into three distinct muscles (Fig. 19). As described by Jenkins and Tanner (1968), the M. stemohyoideus I originates from the medial surface of the scapula and the most anterior part of the clavicle and in- serts on the distal two-thirds of the anterior cornu. This muscle may be the M. ster- nothyroideus of other workers. The M. ster- lohyoideus II originates from the anterola- teral surface of the sternum and inserts onto the posterodorsal surface of the basihyoid. The M. stemohyoideus III is separate for its entire length, with an origin from the ven- tral surface of the anterior third of the ster- num and an insertion on the dorsal surface of the most enlarged area of the posterior comu. In the agamid Chlamydorsaurus, the M. stemohyoideus has a large origin from the sternum immediately deep to that of the M. omohyoideus. It expands and thins as it ex- tends anteriorly to insert on the inner side of the ceratobranchial ventral to the M. omo- hyoideus. In Uromastyx the origin is from both the sternum and the coracoid. Nash and Tanner (1970) describe a super- ficial and a deep layer of this muscle in the skink Eumeces. The larger ventral or super- ficial layer originates from the posterior and ventral surfaces of the ceratobranchial I and ihedial to the corpus and inserts on the inter- clavicle with the M. stemocleidomastoideus, trapezius, and depressor mandibularis, and with the constrictor colli on the posterior and ventral surfaces of the anterior comu. The dorsal or deep layer originates on the inter- clavicle and inserts on the posterior border of both anterior and posterior comua. In Dibamus, the M. stemohyoideus is a large strap originating from the sternum and coracoid and inserting on the distal tip of the ceratobranchial (Gasc 1968). In the teiids Ameiva and Cnemidophorus, the M. stemohyoideus is broad, with an ori- gin on the sternum and insertions on both the posterior and anterior cornua and the basihyoid. In snakes the M. stemohyoideus is found as a separate muscle only in the Typhlopidae and Leptotyphlopidae. Its origin here is deep to the muscles on the linea alba. The fibers pass anteriorly to insert on the hyoid, usually on the entire posterior edge of each comu. 12. M. stemothyroideus In the turtle Trionyx the origin of the M. stemothyroideus is on the anterior border of the stemum. The muscles extend anteriorly to insert on the ventrolateral border of the posterior part of the basihyoid. In lizards, the M. stemothyroideus nor- mally has an origin on the anteromedial por- tion of the sternum, from which it extends anteriorly to insert along the length of the second ceratobranchials. This situation exists in Hemidactylus, Mabuia, Cabrita, Anolis, Calotes, and the iguanine lizards. In Cha- maeleo the M. stemothyroideus extends later- ally to insert on the distal end of the cerato- branchial. In Varanus the M. ster- nothyroideus lies dorsal to the Mm. omo- hyoideus and stemohyoideus. It originates as a thin sheet from the anteromedial half of the sternum and inserts on the anterior half of the proximal piece of the second ceratobranchial. In the iguanine lizards the most medial series of fibers of the M. stemohyoideus com- plex, the M. stemothyroideus, may be sepa- rated from the other members of this group by their different origins and insertions. The origin consists of a small area of both the in- terclavicle and sternum. These fibers pass an- teriorly and parallel to the trachea to insert on the basihyoid. Along its length this muscle is difficult to separate from the more lateral M. stemohyoideus, except in Opiums and Chalarodon, in which both muscles are free and separated along their entire length. 320 Great Basin Naturalist Vol. 42, No. 3 The M. stemothyroideus of Phrynosoma was previously described by Jenkins and Tan- ner (1968) as the M. sternohyoideus I. 13. M. costocutaneous superior Because the shoulder girdle of snakes has been lost, the M. omohyoideus, ster- nohyoideus, and stemothyroideus cannot be identified. Therefore these muscles will be discussed here under the name M. costocu- taneous superior. In some snakes it is possible tentatively to identify the homologs of these three muscles. For example, in the Typhlopidae and the Leptotyphlopidae, the M. sternohyoideus is a distinct mass of fibers that arise from the ven- tral scales and adjacent rows of lateral scales and the ribs, extending anteriorly to the hyoid and surroimding muscles. In Typhlops, Leptotyphlops, Rhinopliis, Cylindrophis, and Achrochordus, the anteriormost fibers of the complex extend to originate on the mandible and overlay the hyoid while having no con- nection with it. In Cylindrophis the fibers originate on the posterior or medial edge of the M. constrictor colli. In the anomalepidid snakes the insertion is on the posteromedial border of the basihyoid and second cerato- branchial. The insertion also extends to the base of the lingual process in most specimens. In Agkistrodon, Bothrops, and Crotalus, the insertion is most extensive on the median raphe and lingual process. Sondhi (1958) describes three specific mus- cles present in Matrix (Xenochrophis) that are probably homologous to the Mm. omo- hyoideus, sternohyoideus, and ster- nothyroideus. The omohyoid portion arises from the skin on the ventrolateral part of the neck and then extends obliquely anteriorly to insert on the ventrolateral aspect of the basi- hyoid. In Atretium, this muscle has a cut- aneous origin and inserts on the second ce- ratobranchial. The second muscle, absent in Atretium but possibly the M. sternohyoideus, originates from the skin in the ventrolateral region of the neck posterior to the M. omo- hyoideus and passes anteriorly to close prox- imity with the latter to insert on the outer border of the basihyoid. The sternothyroid part of this complex lies in the midline of the neck over the ventral surface of the basi- hyoid, with its origin on the midventral por- tion of the cervical skin. The muscle inserts on the medial border of the basihyoid. In Atretium the sternothyroid portion of the complex has its origin from the second ce- ratobranchial, with some fibers intertwining with their opposite member at the midline. 14. M. neurocostomandibularis According to Langebartel (1968), the M. neurocostomandibularis is present in all snakes except the Anomalepididae. In most snakes it is a broad sheet forming part of the M. neurocostomandibularis complex, but in some it is separate and narrow. It covers a large area of the head and in some is partially overlain by the Mm. constrictor colli and cos- tocutaneus superior. Its origin is on the den- tary, from which it proceeds posteriorly to insert variously on the hyoid apparatus. The muscles of Python sebae (Frazzetta 1966) and Boa constrictor (Gibson 1966) that are innervated by the hypoglossal nerves form a single muscular complex, the M. neurocostomandibularis, and correspond roughly to the M. geniohyoideus of other reptiles. The complex extends between the mandibles and the second ceratobranchials. In both Boa and Python the origin is on the lower jaw and the insertion on the posterior part of the second ceratobranchial. In Natrix (Xenochrophis), Sondhi (1958) de- scribes the M. neurocostomandibularis as probably the M. geniolateralis because the latter muscle receives a branch from the hy- poglossal nerve. Langebartel (1968) consid- ered this muscle to be the M. ceratomandibu- laris as designated by Richter (1933). The proper identity of this muscle in the typhlo- pids, leptotyphloids, and anomalepidids is un- known to us. According to Langebartel (1968), the M. ceratomandibularis in snakes arises from the dentary and itiserts on the an- terior part of the hyoid and the tendinous in- scription in the M. neurocostomandibularis. Varkey (1979) describes the M. neurocosto- mandibularis as being very complex and hav- ing three separate heads in Nerodia. It is a wide flat muscle sheathing the neck and most of the lower jaw. One origin (the vertebral head) is on the apponeurosis of the dorsal September 1982 Tanner, Avery: Buccal Floor of Reptiles 321 midline neck region. It passes under the con- strictor colli to insert on the midline raphe. The costal head originates by narrow slips from the first seven anterior ribs and inserts on the midline raphe with the previous slips. The third or hyoid head has a double origin from the midventral raphe just median to the hyoid comua. This branch is called M. trans- versalis branchialis by Langebartel (1968). It inserts on the origin of the other heads on the midline raphe. 15. M. transversalis branchialis The M. transversalis branchialis appears variably and erratically in the families of snakes with the exception of the Anoma- lepididae, Typhlopidae, and Leptotyphlo- pidae, in which it is universally absent. When present, this muscle arises from somewhere on the second ceratobranchial. In Rhinophis, the origin is on the medial edge, whereas in Cylindrophis it originates on the anterior two-thirds. In Anilius the entire length of the cartilage is involved. The insertion of this muscle is usually on the median raphe, although in some snakes it is inserted on the fascia covering the M. cos- tocutaneous superior. In Nerodia, Varkey (1979) describes the M. transversus branchialis as originating on the midline raphe just anterior to M. inter- mandibularis's anterior. It passes anterolate- rally to insert broadly on the mucosa of the angulo-splenial articulation and narrowly on the lateral sublingual gland. Varkey indicates his usage of this muscle name is as in Albright and Nelson (1959), Cowan and Hick (1951), and Weaver (1965). Langebartel (1968) calls this muscle the dilator of the sublingual gland, using the name M. transversalis bran- chialis for a branch of what Varkey calls the M. neurocostomandibularis. 16. M. hyotrachealis In most snakes the M. hyotrachealis arises from the second ceratobranchial, but in Liotyphiops and the leptotyphiopids the fi- bers are tied by connective tissue on the ven- trolateral surface of the lining of the buccal floor. In the typhlopids the fibers originate in connective tissue on the hypaxial trunk mus- cles. In other snakes the M. hyotrachealis originates on the lateral edge of the second ceratobranchial. In Rhinophis the origin is at the anterior quarter of the medial edge, while in Cylindrophis maculatus and C. rufus the origin is from the lateral edge about half- way down the ceratobranchial. In the boids it originates on the posterior half of the carti- lage. In Tropidophis the origin is deep from the raphe of the M. neurocostomandibularis. In colubrids, viperids, and elaphids the origin varies extensively. In Heterodon and Pseu- daspis the origin varies extensively. In Heter- odon and Pseudaspis the origin is from the rib cage, while in Agkistrodon it may be ei- ther the rib cage or hyoid, indicating a split origin. In Vipera aspis, Edgeworth (1935) de- scribes one head oiF the M. hyotrachealis as lying dorsal to the rib cage while the lateral head attaches to the hyoid. In Cerastes the single head originates from the ventral lining of the buccal floor. The insertion of the M. hypotrachealis is normally from the trachea of the laryngeal- tracheal area, dorsal and anterior to the in- sertion of the M. geniotrachealis. In some genera {Typhlops, Amblycephalus, Xeno- peltis, and Agkistrodon piscivorus) the in- sertion is on the ventral portion of the M. geniolateralis. In Boa cookii, Notechis, and others the M. hypotrachealis has a split in- sertion with attachments on dorsal and ven- tral sides of the geniotrachealis. Varkey (1979) describes Nerodia' s. hyotra- chealis as thin and narrow and of a double origin. One head is just anterior to a trans- verse tendinous inscription of the M. neuro- costomandibularis. The second or median head is from the lateral edge of the hyoid comua. The heads join and insert on the lar- ynx and trachea anterior to the insertion of the geniotrachealis. V. Buccal Floor Muscles Not Associated with the Hyoid Apparatus The homologies of a number of the repti- lian throat muscles not connected with the hyoid are unclear. We will present the most widely used terminology and present syno- nyms only when two or more names have had wide usage for the same muscle. Al- though the following muscles are not directly attached to the hyoid apparatus, they have a 322 Great Basin Naturalist Vol. 42, No. 3 close functional relationship and are there- fore included (Figs. 18, 19, 20). 1. M. constrictor superficialis The M. constrictor superficialis is found in Trionyx as a superficial muscular sheet lying ventral to the anterior region of the neck. It arises as a narrow slip from the skin covering the side of the neck and broadens to insert on the gular septum. In Gavialis it originates on the skin overlaying the angle of the jaw, sur- roimds the neck, and extends obliquely to in- sert on the gular septum. In other reptiles, such as iguanid hzards, this muscle is probably homologous to much of the Mm. constrictor colli and inter- mandibularis posterior of Avery and Tanner (1971). 2. M. constrictor colli The M. constrictor colli is an extensive su- perficial muscular sheet, originating on the middorsal aponeurosis of the neck and ex- tending ventrad to insert on the posterior part of the midventral raphe or gular septum. In Sphenodon it forms a broad, thin, super- ficial sheet that completely encases the neck (Rieppel 1978, Fig. 1). It ensheathes the en- tire neck in Chehdina and Deirochelys, but in Lissemys the neck is only partly covered. The muscle arises from the dorsal fascia and inserts on the median raphe. It is continuous anteriorly with the M. intermandibularis. Sondhi (1958) lists this muscle as present in Trionyx and Gavialis, and Gnanamuthu (1937) recognized it in Crocodylus and Tri- onyx. In the Testudines and Crocodylia the M. constrictor colli is not attached to the hyoid, but has an insertion on the gular septum. This muscle covers most of the lateral sur- face of the neck in Amblyrhynchus, Chalaro- don, Cyclura, Iguana, and Saiiromalus. It is much less extensive in Brachylophus, Con- olophus, Ctenosaura, Dipsosaunis, Opiums, Crotaphytus, and Phrynosoma. In Chamaeleo the M. mylohyoideus poste- rior of Mivart (1870) corresponds to the M. constrictor colli. It originates on the occipital crest and inserts on the median raphe. In the skink Eumeces the M. constrictor colli is a very broad sheet originating from the middorsal tympanic fascial and inserting on the median raphe. It covers most of the neck from the angle of the jaw to the interclavicle. In the teiids Cnemidophorus and Ameiva, the muscle is as in Eumeces, but the anterior border interdigitates with the posterior bor- der of the M. cervicomandibularis. Gnana- muthu (1937) figures this muscle to be in Hemidactylus, Mabuia, Cabrita, Anolis, Ca- lotes, and Draco. In snakes the M. constrictor colli appears erratically and is not constant in form within a single genus as indicated by Python. The muscle is normally broad and envelops the angle of the jaw with an insertion on the midventral raphe or hyoid. In some species of Python it appears to be absent. The M. con- strictor colli is found in all families of snakes except in the Uropeltidae where it has not been recognized. 3. M. mylohyoideus anterior The superficial M. mylohyoideus anterior is generally located ventrally beneath the rami of the lower jaw anterior to the M. con- strictor colli. It takes its origin from the ante- rior part of the mandible and inserts on the gular septum. In Sphenodon the M. mylohyoideus (M. in- termandibularis of Rieppel 1978) forms a single large muscular sheet, but in lizards it is differentiated into three sets; the Mm. mylo- hyoideus anterior superficialis ( = M. inter- mandibularis anterior superficialis), mylo- hyoideus anterior principalis ( = M. intermandibularis anterior profundus), and mylohyoideus anterior profundus ( = M. inter- mandibularis posterior). In some forms, such as Cabrita, Anolis, Hemidactylus, and Mabuia, some fibers of the M. mylohyoideus anterior originate deep on the medial surface of the mandible and others originate super- ficially on the M. geniohyoideus. As the fi- bers of the two muscles cross, they break into numerous strips and interdigitate (Figs. 18, 19, and 20). In turtles the M. mylohyoideus is simpler. In Trionyx it consists of a single M. mylo- hyoideus anterior profundus that originates September 1982 Tanner, Avery: Buccal Floor of Reptiles 323 ventral to the M. geniohyoideus from the ventral aspect of the mandible and passes medially to insert on the gular septum. In Lissemys the M. mylohyoideus anterior forms as two muscles, with the M. mylohyoideus anterior profundus being identical to that of Trionyx. The M. mylohyoideus anterior prin- cipalis is a broad sheet originating on the mandible and inserting on the gular septum. In Chelodina and Deirochelys the inter- mandibular series is simple and undivided, originating on the inner surface of the man- dible and inserting on the median raphe. In Chelodina the anterior quarter of the fibers do not insert into each other as in Deiro- chelys, but are separated by fascia. Some age variation in this muscle can be seen in Crocodylus: in adults the M. mylo- hyoideus anterior is not distinguishable as a separate muscle, but there are two sheets in the juvenile representing the M. mylo- hyoideus anterior and mylohyoideus posteri- or. In adult Alligator a single transverse sheet is present (Mm. intermaxillaris and sphincter colli), and in Gavialis the one sheet (M. mylohyoideus anterior principalis) is prob- ably homologous to both the Mm. mylo- hyoideus anterior and mylohyoideus posteri or. In Gavialis this muscular sheet occupies almost the entire anterior part of the ventral inter-ramal area of the neck, originating on the inner side of the mandible and inserting on the gular septum. The M. mylohyoideus anterior superficialis exhibits several variations. In Mabuia and Anolis the fibers extend anteriorly, over- lapping the M. mylohyoideus anterior princi- palis either medially or laterally. In the Cha- maeleonidae and Agamidae the Mm. mylohyoideus anterior principalis and mylo- hyoideus anterior profundus occur together in the fonn of a double sheet, which we have concluded is a variation of the M. mylo- hyoideus anterior. In Varanus, Sondhi (1958) indicates that the muscle extends transversely from the mental groove to the M. gen- ioglossus portio major. There are three sets of fibers listed, including a broad M. mentalis superficialis, that originate ventrally, whereas the narrow M. mentalis profundus anterior and the M. mentalis profimdus posterior orig inate dorsally. All tliree bundles insert in the lining of the buccal floor. These muscles do not appear to be homologous to the muscular complex we have seen in iguanids and desig- nated the M. mylohyoideus anterior. In Amblyrhynchus, Brachylophus, Chalaro- don, Conolophus, Ctenosaura, Cyclura, Dip- sosaurus. Iguana, Opiums, and Sauromalus two distinct groups of muscle fibers are found. The M. intermandibularis anterior are deep fibers that originate as a tendon from the coronoid and splenial bones and extend medially on the ventral surface to join at the median raphe, where they interdigitate with about five bundles of the M. geniohyoideus. A small bundle of fibers also extends from the origin to insert on the connective tissue cap- sule of the sublingual gland. In Iguana this muscle forms the bulk of the large dewlap. The most superficial group of fibers (M. in- termandibularis anterior superficialis) is small and narrow, with an origin from the oral membrane and from the anterior part of the M. intermandibularis anterior profundus. The muscle fibers pass obliquely posteriorly to in- sert on the median raphe. In Iguana and Dip- sosaurus this superficial group is greatly re- duced in size. In snakes the synonymy of the throat mus- culature is not well established. For this rea- son we follow rather closely the studies of Langebartel (1968) and Sondhi (1958). The anteriormost set of transverse fibers (M. in- termandibularis anterior) is absent in Anilius and Xenopeltis, but is represented by a ten- don in Rhinophis. In the anomalepidids, typhlopids, and leptotyphlopids this muscle is broad and may actually represent several muscles. In the latter families one or more muscle groups may originate on the medial surface of the dentary. In the colubrids, vipe- rids, and elapids a single muscle is large but separated into two parts. The longer and thicker anterior one originates on the medial surface of the tip of the dentary and medially to the fibrous inter-ramal pad. The second (posterior) part extends obliquely from the same origin to insert on the ventral raphe. The M. mylohyoideus anterior in Matrix {Xe- nochrophis) is probably represented by three muscles: Mm. intermaxillaris, mentalis pro- fimdus anterior, and mentalis profundus pos- terior. The M. intermaxillaris originates from the ventrolateral border of the dentary and passes obliquely posteriorly to insert on the 324 Great Basin Naturalist Vol. 42, No; 3 mental groove. The remaining pair of bun- dles originate from the mental groove and ex- tend obliquely caudad to insert adjacent to each other on the lining of the buccal floor. The intermandibularis anterior of Nerodia is described as having two separate parts. The M. intermandibularis anterior pars mucosalis has two portions. The first is a small triangular anterior portion that origi- nates on the midventral raphe of the lower jaw and buccal membrane fascia. It inserts on the ventromedial surface of the anterior tip of the dentary and the ligament attached to it. The much stouter posterior slip originates from the midventral raphe of the lower jaw and fascia surrounding the tongue sheath just posterior to the insertion of the anterior slip. The fibers pass anterolaterally to insert on the ventromedial surface of the dentary im- mediately posterior to the insertion of the an- terior slip. The second part (M. intermandibularis an- terior pars glandularis) originates on the mid- ventral raphe of the lower jaw of the fibrous inter-ramal pad. The fibers pass post- erolaterally to insert on the ventrolateral side of the sublingual gland at its posterior end. A small number of fibers insert on oral mucosa posterior to the gland. 4. M. mylohyoideus posterior The M. mylohyoideus posterior is a trans- verse muscle situated posterior to the M. mylohyoideus anterior. The M. mylohyoideus posterior [M. mylo- hyoideus anterior principalis of Sondhi (1958)] of Trionyx and Lissemys originates from the border of the mandible, where it forms a broad, thick sheet muscle. It extends medially to insert on the gular septum. In Alligator, Crocodylus, and Gavialis the M. mylohyoideus posterior is represented by a thin sheet that combines into one muscle, the M. mylohyoideus anterior and M. mylo- hyoideus posterior. In some lizards {Mahuia and Cabrita) these muscular sheets are continuous, but show a small division between them. Sondhi (1958) reports that in Varanus they are differen- tiated into two muscles (Mm. mylohyoideus anterior superficialis and mylohyoideus ante- rior principalis) that are disposed one behind the other, both originating on the lateral sur- face of the mandible and inserting on the gu- lar septum. In the iguanine lizards, an anterior and a posterior sheet of muscle fibers (M. inter- mandibularis posterior) form the M. mylo- hyoideus posterior. The anterior sheet is broad and thin, with an origin from the later- al surface of the mandible. The fibers pass medially on each side to insert with their op- posite members at the median raphe. The posterior bundle of fibers (about one-quarter of the posteriormost fibers) originate from the lateral surface of the mandible beginning at the midpoint of the retroarticular process and insert on the linea alba. In most iguanines the M. mylohyoideus posterior exhibits a primitive condition by being continuous with the M. constrictor col- li, from which it can be delineated by a natu- ral separation along the entire border only in Conolophus and Ctenosaura. In Cyclura and Sauromalus this separation is present only in the medial third of their common border. In Amblyrhynchus, Brachylophus, Chalarodon, Dipsosaurus, Iguana, and Opiums the two muscles are continuous along their entire border. In Crotaphytus the Mm. mylohyoidei ante- rior and posterior form one continuous sheet with no separation between them. In Phryno- soma the M. mylohyoideus posterior is sepa- rated from the anterior, but is continuous posteriorly with the M. constrictor colli, from which it can be separated only with great care. In Eumeces the position of M. mylo- hyoideus posterior is similar to that of the iguanid lizards, with both anterior and poste- rior muscles being separated. In the teiid Ameiva the M. mylohyoideus posterior originates on the medial surface of the dentary and immediately breaks into nine separate divisions that interdigitate with slips of the M. geniohyoideus. It inserts on the midventral raphe just posterior to the M. mylohyoideus anterior (Fig. 17). In Cnemido- phortis the muscle is as above, except that there are only five divisions instead of the nine in Ameiva. The Mm. mylohyoideus pos- terior and constrictor colli are continuous for their entire border in Shinisaurus, but widely separated in Xenosaiirus. September 1982 Tanner, Avery: Buccal Floor of Reptiles 325 All snakes except one colubrid {Amhlyce- phahis kuangtunensis) possess a M. inylo- hyoideus posterior (Langebartel 1968). It lies in the same position as the M. inter- mandibularis posterior of Langebartel (1968), with the former having its origins on the mandible and insertion on the gular septum. In the colubrid Achrochordiis it is very broad, attaching to the middle region of the man- dible. In Haplopeltura boa it is attached lat- eral to the external adductor muscle of the lower jaw. The fibers are not attached to the mandible, but cross their opposite members at the midventral raphe and interdigitate, eventually attaching to the opposite man- dible. M. mylohyoideus inserts on the lingual process in the hydrophid Aipysumrus. The second bundle of fibers (M. inter- mandibularis posterior superficialis) is small and restricted in some snakes with the paral- lel type of hyoids. Its occurence is sporadic in colubrids, and it is absent in most poisonous snakes, including the hydrophids. It may be replaced by a tendon that originates from the posterior part of the lower jaw and inserts on the gular septum. Sondhi (1958) states that the M. mylohyoideus posterior is an extreme- ly broad muscle sheet, lying immediately pos- terior to the M. mylohyoideus anterior and occupying the posterior region of the neck in Matrix piscator. He further states that this muscle originates on the dorsolateral surface of the anterior cervical vertebrae and extends ventrally to insert on the posterior part of the gular septimi. Langebartel (1968) describes two muscles, a ventral sheet taking its origin from the anterior part of the mandible and extending obliquely anteriorly over the body of the tongue to insert on the gular septum, and a dorsal sheet deep and dorsal to the M. geniohyoideus, with an origin from the man- dible with the ventral sheet; the dorsal sheet extends obliquely anteriorly also to insert on the gular septum. The M. ceratomandibularis of Langebartel (1968) occurs in most snakes, although with considerable variation. Be- cause of its location, we include it as a syno- nym of M. mylohyoideus posterior, even though we are aware that most homologies must yet be proven by careful embryonic study. The intermandibularis posterior of Nerodia is described by Varkey (1979) as having two slips. The M. intermandibularis posterior pars anterior is the largest of the ventral con- strictors originating on the midventral raphe of the lower jaw ventral to the origin of the posterior slip of the intermandibular anterior I and the transversalis branchialis. The origin is broad and thin, passing caudolaterally to form a stout band to insert on the ventrome- dial surface of the bone at the distal end of the mandibular fossa. The second slip, which Varkey calls the M. intermandibularis posterior pars posterior, is a thin, flat, triangular sheet of muscle that originates on the midventral raphe posterior to the origin of the intermandibularis posteri- or I and the transversalis branchialis and the anterior tip of the hyoid cornua. It passes ventral to the M. neurocostomandibularis for most of its length. This insertion is just poste- rior to the insertion of the pars anterior on the ventrolateral surface of the bone at the level of the proximal end of the mandibular fossa. 5. M. mandibulotrachealis The M. mandibulotrachealis of Varaniis has been described by Sondhi (1958) as a deli- cate muscle arising from the anteroventral part of the mandible and extending posteri- orly to divide into two parts. The dorsal part passes posteriorly dorsal to the tongue to in- sert on the lateral side of the trachea. The ventral part of the muscle extends posteriorly to fan out over the buccal floor near the in- sertion of the M. genioceratoideus, with an insertion on the ventral lining of the buccal floor. In Natrix (Xenochrophis), Sondhi (1958) found the origin to be similar to that of Va- ranus, with a medial bundle inserting on the trachea and a lateral bundle attaching to the lining of the buccal floor. It has not been re- ported for other genera. This muscle is reported by Varkey (1979) for Nerodia as the M. geniotrachialis. It is a stout band of muscle that parallels the gen- ioglossus. It originates at the anterior end of the dentary dorsal to the origin of the lateral genioglossus. It passes posteromedially to the tongue sheath and inserts on the ventral and ventrolateral surfaces of the first 14 tracheal rings. 326 Great Basin Naturalist Vol. 42, No. 3 6. M. neuromandibularis The M. neuromandibularis has been de- scribed in detail by Sondhi (1958) for Matrix (Xenochrophis) and Varanus. In the latter it probably corresponds to the M. gen- iolateralis. Sondhi has described M. neuro- mandibularis as originating from the dorsola- teral border of the middorsal aponeurosis and extending a short distance anterior along the dorsal side of the neck. The fibers divide into three sets, which pass into a common tendon inserting on the inner ventral surface of the skin. In Natrix (Xenochrophis) the origin is similar to that in Varanus. The insertion is on the ventromedial side of the posterior half of the mandible. In some snakes the M. neuromandibularis arises from an aponeurosis at the middorsal line. In the anomalepidids, typhlopids, lepto- typhlopids, uropeltids, and aniliids it inserts on the lov^^er jaw. In the Xenopeltidae, Boidae, and other families it inserts on the raphe in common with the Mm. ceratoman- dibularis and costomandibularis. 7. M. costomandibularis The M. costomandibularis has been de- scribed only for some snakes in which its ori- gin is either from the cartilaginous ribs or the rib cage. In Thamnophis a medial slip origi- nates from the peripheral surface of the lin- ing of the pharyngeal floor and inserts on the common tendinous inscription of the M. neurocostomandibularis. In Cylindrophis rufiis the insertion is on the second cerato- branchial as well as on the mandible. 8. M. constrictor pharyngis The M. constrictor pharyngis of Croco- dylus and Gavialis is a deeply laid transverse strap apparently restricted to the Crocodilia. Its origin is from the lateral surface of the cervical vertebrae and its insertion medial on the gular septum. 9. M. obliquus abdominis internus Langebartel (1968) describes the M. ob- liquus abdominis internus as a trunk muscle of snakes, with an origin on the medial face of the ribs and an insertion on the linea alba. 10. M. transverse abdominis Langebartel (1968) has also described the M. transverse abdominis as restricted to snakes and lying on the deep surface of the M. obliquus abdominis internus, with an ori- gin on the medial face of the ribs. After ex- tending posteriorly and medially, it inserts on the linea alba. VI. The Tongue: External Morphology The tongue of reptiles has been in- vestigated by many workers, some of which are as follows: Graper (1932), Nonoyama (1936), Gnanamuthu (1937), Oelrich (1956), Sondhi (1958), Avery and Tanner (1971), and Kroll (1973). Winokur (1974) published a ma- jor study on the adaptive modification of the buccal mucosae in turtles. His study is con- cerned not only with the tongue, but also with the glands found in the buccal area. Tongues in turtles vary in size and com- plexity (Fig. 22). Winokur states that Terrestrial herbivores {Gopherus, Testudo, and other tor- toises) have the best developed mucous glands, whereas aquatic carnivores (Chelydra and Chelus) have few or no mucous glands. In both Chelydra and Trionyx (Fig. 22 A, B) the tongue is without papillae or complex glands and is nonprotrusible, a characteristic of carnivorous chelonians. In Trionyx the short, rounded tongue is dorso-ventrally flat- tened and contains just a base and body. The base is formed from two posterior limbs that they enclose. Each basal portion extends an- teromedially to unite in the tongue. Posterior to the tongue and glottis, the buccal-phary- ngeal floor has numerous filiform papillae that Girgis (1961) has shown to have a res- piratory function. The tongue of some, such as Chehis, has been developed as a lure in food-getting: the open mouth exposes a wormlike tongue structure to intice unsus- pecting prey into the mouth. In contrast, the terrestrial herbivorous che- lonians (Tortoises; Fig. 22b) have a much September 1982 Tanner, Avery: Buccal Floor of Reptiles 327 larger lingual pad, which is glandular, fleshy, and somewhat protrusible. Tortoises have profuse Ungual mucous glands on and be- tween the lingual papillae as well as muscles capable of some lingual protrusion. They generally lack papillae posterior to the tongue. Winokur (pers. comm.) considers Derrna- teinys (Fig. 22a) to be a special case. The tongue of this aquatic herbivore shows one end of the spectrum of buccal complexity in aquatic chelonians. Figures 22 and 23 illus- trate the extremes seen between the tongues of aquatic carnivorous and terrestrial herbi- vorous chelonians. The tongue of Derma- temys, although proportionately smaller than that of terrestrial Gophems, shows an ex- treme condition of buccal papillation, but one that is quite different from that of terres- trial herbivorous tortoises. Between these ex tremes are the majority of chelonians, such as Pseudemys, which tend toward omnivorousness. The tongues of Alligator, Crocodylus, and Gavialis lack any specific areas identifiable as base, body, or apex. The tongue is a mass of tissue between the mandibular symphysis and glottis attached to the lining of the buc- cal floor except at its anterior tip. It can be elevated and depressed, but not protruded. Sauromalus (Fig. 24) and Brachylophus (Fig. 25) show generalized lizard tongues, with their extensive papillation and lateral extensions on each side of the glottis. Such tongues are protrusible and obviously serve a masticatory function. In Amblyrhynchus, Brachylophus, Conolophus, Ctenosaura, Cycl- ura, Dipsosaurus, Iguana, and Sauromalus the tongue is well developed and large. In the above genera it is cleft anteriorly, with the most anterior tips lacking papillae. There is a smooth pad ventral to the tips (Fig. 24). In the teiid Ameiva the tongue is rounded and slightly notched posteriorly and covered by a lingual sheath. It bears a deep terminal notch anteriorly that separates the tapering elongate terminal prongs. A lingual sheath is absent in Cnemidophorus and other macro- teiids. The tongue of Cnemidophorus (Fig. 26) represents a moderate advancement in the development of flexibility, and Cha- maeleo (Fig. 27) is a highly specialized free tongue. Fig. 22. Tongue size as indicated in: A, Dermateys mawi (UU 9845), above; B, Gophems (UU 5961), below. It should be noted that the glottis is moved caudad as the tongue increases in size. Photographs provided by Robert M. Winokur. 328 Great Basin Naturalist Vol. 42, No. 3 PhF Fig. 23. Tongue of Trionyx spiniferus: A, showing its position in relation to the glottis and pharynx area with its filamentous papillae; B, Chelydra serpentina showing the nonpapillated pharynx. In Lanthanotus the tongue is deeply in- cised terminally, forming two tapering prongs. The anterior half of the tongue is elongate, narrow, smooth, and elastic, where- as the posterior half is wide and covered with papillae. In Shinisaurus the tongue is similar, but the posterior half is more triangular and the terminal prongs are not as well devel- oped. Heloderma has a similar tongue but with proportionately longer terminal prongs than in the latter. The tongue of Varanus is elongated and protrusible, terminating in a forked tip ante- riorly. The entire median part of the buccal floor is occupied by its mass. Posteriorly it extends as a bifurcated portion on each side of the glottis and esophagus and into the neck proper. Sondhi (1958) considers the tongue to be divisible into three parts: the base which is bifurcated; the body, formed by the union of two basal masses of muscle; and lastly the apex, consisting of a pair of prongs. Each muscular mass forming the basal branch of the tongue arises on the distal end of the second ceratobranchial as a slender longitudi- nal M. hypoglossus, which extends along the ventrolateral surface of each ceratobranchial to pass obliquely to the dorsolateral side of the neck. This muscle eventually occupies a midventral position, with the middle of its basal branch lying ventromedial to the point of articulation between the distal and pro- ximal portions of the second ceratobranchial. Its anterior portion lies ventrolateral to the proximal piece of the anterior cornu at the point of articulation with the basihyoid. As the two basal branches of the tongue approach, they become thick and sub- cylindrical and eventually lie dorsal to the basihyoid and ventral to the portiones pro- September 1982 Tanner, Avery: Buccal Floor of Reptiles 329 Fig. 24. Tongue of Sauwmahis obestis: A. outline of dorsal view; B, ventral view showing the smooth pads sur- rounding the tips. (Dorsal surface as in Brachijhphiis, Fig. 25). ximales of the anterior cornua. At their ante- rior extremes the two branches are enclosed in a Ungual sheath, where they unite to form the body of the tongue. The body is enclosed by the lingual sheath and occupies the medial area of the buccal floor. Ventrally the ante- rior end of the lingual process lies inside the lingual sheath and opposite the glottis. Also, ventrally the two handlelike pieces of the portiones proximales overlap medially to cover the body of the tongue. The apex of the tongue consists of a pair of prongs. rounded, thick at the base, and tapering to pointed ends anteriorly. We recognize at least three types of sau- rian tongues. First, in the generalized tongue, seen in such forms as Sauromalus and Coleo- nyx, the dorsal surface is papillate and highly glandular; although the tip is divided, it is not extended into a pair of elongated prongs. Second, an elongate, narrow tongue with a pair of elongate prongs occurs in such groups as the teiids and varanids. In these lizards with deeply incised tips, the tongue is narrow 330 Great Basin Naturalist Vol. 42, No. 3 and glandular and serves not only the pur- pose of mastication, but also functions as a sensory organ. Sondhi (1958) implies that such tongues are closely related anatomically to the tongues of snakes, and he compares the tongue of the natricine snakes to that of Va- ranus. Third, an entirely different tongue is found in Chamaeleo. Instead of a further de- velopment of the tip as in Varanus, the cha- maeleonids have developed a blunt end with a highly glandular anterodorsal surface used in capturing and ingesting food. In snakes the tongue has developed a greater bifurcation with filamentous lateral projections on each fork. Such tongues are sheathed at their base and function as a sen- sory rather than a masticatory or food-getting Fig. 25. Tongue of Brachylophus showing the size and nature of the tongue papillae (TP), and the reti- culated, ridged nature of the tissue (B) extending poste- rior to the glottis (G). organ. Our imderstanding of lingual struc- tures and the associated buccal mucosae, however, is still sketchy and much com- parative study must be done before an ade- quate understanding of their anatomy is available. In Matrix (Xenochrophis) Sondhi (1958) also describes the tongue as having three parts, with the basal branches lying parallel on each side of the midlongitudinal Hue ventral to the trachea. Each branch passes anterior to the second ceratobranchial ventromedially. As they approach the dorsal part of the basi- hyoid, the two branches unite to form the body of the tongue, which is elongated and compressed dorsoventrally. In the retracted position the tongue is almost entirely encased by the lingual sheath dorsal to the basihyoid and lingual process and ventral to the trachea. The apex of the tongue is broad at the base but tapers anteriorly. The tongue has a variety of forms, sizes, and functions in reptiles. In some aquatic tur- tles it is a small pad rather tightly applied to the floor of the anterior part of the mouth. Such tongues are nonprotrusible and actually have a very limited ability to move. In most chelonians, except for some aquatic turtles and crocodilians, the tongue is more than a pad and serves many useful functions. In some chelonians {Gopherus, Fig. 22b), most lizards (iguanids and agamids for example. Fig. 24), and in the more primitive Spheno- dontidae the tongue may serve a masticatory function. It is a "food-getting" organ in the "free"-tongued Chamaeleonidae and has a sensory function in snakes and some lizards. As noted above, the degree of flexibility in the tongues of reptiles varies from little to considerable movement. Because tongues in most reptiles (except snakes) are associated with feeding, that is, ingestion, their anatomy and perhaps the degree of flexibility is de- pendent on adaptive change to meet such activities. In the Sphenodontidae and Chamaeleo- nidae the extremity is very blunt (Fig. 27). The Chamaeleo tongue and its associated muscles and other tissues may be as long or longer than the body when fully extended. A broad fleshy tongue with smooth and papil- late areas is seen in the gekkonids and igua- nids (Figs. 24, 25). The Testudines and September 1982 Tanner, Avery: Buccal Floor of Reptiles 331 Fig. 26. Dorsolateral view of Cnemidophorus tigris (BYU 17366) showing the forked tongue and the narrow papil- late body of the tongue (B), floor of the mouth (M), cut muscles. Crocodilla may have small pads with little movement or, as in those such as Gopherus, the tongue is larger, fleshy, and closely tied to the buccal floor and has varying protnisibility. The highly flexible and protrusible tongue of snakes has become an elongate, slender, sensory organ. In this form it has changed to an entirely different organ than that of most other reptiles, in which the tongue is an or- gan lying on the buccal floor. In its normal position it is sheathed, with little or none of it visible on the buccal floor. Also, the open- ing of the tongue sheath has moved anterior so as to lie just posterior to the mental syn- thesis, with the glottis immediately posterior to the sheath opening. Although ophidian tongues are structurally and functionally quite different from those of most other rep- tiles, they are nonetheless developed phy- logenetically from the same basic structures. The adaptive changes found in the tongues of reptiles are probably some of the most re- markable to be found, for one organ, in the vertebrate series. VII. Musculature of the Tongue The tongue is associated with musculature of two basic types: (1) extrinsic musculature, which does not contribute to the structure of the tongue itself, and (2) intrinsic muscula- ture, which makes up the lingual structures. 1. Extrinsic musculature In most reptiles the M. geniohyoideus is the primary extrinsic muscle of the tongue. It is paired and arises from the mandibular sym- physis to insert on the external part of the M. hypoglossus, parts of the hyoid apparatus, or the lining of the buccal floor. In Sphenodon it has two extensions, one dorsal and one ventral. In the turtles Trionyx and Lissemys the M. geniohyoideus is undivided and broad. It originates on the mandibular symphysis and extends posteroventrally to insert on the fas- cia of the ventrolateral border of the body of the tongue. The M. geniohyoideus of Alligator takes origin from the mandibular symphysis and di- 332 Great Basin Naturalist Vol. 42, No. 3 Fig. 27. Tongue of Chamaelon brevicarnis: A, lateral view showing position of tongue in mouth cavity; B, ventral view with muscles and other tissues removed to show the tongue and the folded M. hypoglossus; C, tongue removed, lateral view (BYU 12422). vides into medial and lateral bundles. The medial bmidle is narrow and interdigitates with its opposite member to insert on the basihyoid. The lateral bundle is broader and inserts on the tongue. The M. geniohyoideus of Crocodylus arises from the mandible and divides into two lateral bundles, both of which extend posterodorsal to where the me- dian bundle of the M. geniohyoideus inserts on the anterior border of the hyoid. The lat- eral bundle inserts on the anterior and ven- tral border of the anterior comu. In Gavialis the M. geniohyoideus has portiones minor and major, with the portio minor being slen- der and originating with the mandibular sym- physis. It extends caudad to insert on the ven- tral part of the M. hyoglossus. The broader portio major lies lateral to the M. hypo- glossus, takes origin from the mandibular symphysis dorsal to the portio minor, and ex- tends obliquely caudad to a fanlike insertion on the fascia near the middle of the M. hyog- lossus (Sondhi 1958). In Hemidactylus the M. geniohyoideus is well developed, with insertions on the ven- trolateral surface of the tongue and the hyoid comu. In Anolis, Sitana, Calotes, and Draco the M. geniohyoideus fans out to insert on the buccal floor, with the main body attach- ing to the sides of the second comu and the first ceratobranchial. The M. geniohyoideus of Mabuia covers the M. hypoglossus on its lateral surface, whereas in Cabrita it origi- nates on the medial sides of the mandible. The muscle extends posteriorly to insert on the lining of the buccal floor. In the area of the glottis the main bundles of the M. gen- iohyoideus divide into two and insert on the first ceratobranchial on the ventral side of the M. hypoglossus. The M. geniohyoideus of Chamaeleo brevi- cornis consists of two main bundles: the dor- sal one inserts on the buccal floor, the ventral one on the body of the hyoid and the first ce- ratobranchial, lacking any connection with the tongue. The dorsal bundle has three slips September 1982 Tanner, Avery: Buccal Floor of Reptiles 333 that insert (1) on the side of the pouch in the buccal floor where the tongue retracts, (2) on the buccal floor two-thirds the length of the jaw, and (3) after extending obliquely under the second bundle, on tissue lateral to that bundle. In Varanus the M. geniohyoideus, accord- ing to Sondhi (1958), can be divided into two parts. The Mm. geniohyoidei portio minor is very short, extending obliquely post- eromedially to insert on the anterior part of the lining of the buccal floor near the mid- line, and the M. geniohyoideus portio major extends posteriorly for a much longer dis- tance to meet its opposite member at the midline. As the two muscles lie together at the midline ventral to the tongue, each sepa- rates into a dorsal and ventral sheet. Each of these divisions insert on the ventral, lateral, and dorsal sides of the tongue to attach on the fascia of the basal branch of the tongue. In Varanus indicus the main body of the M. geniohyoideus inserts in a fascia in common with the M. stemohyoideus and to the first ceratobranchial, which lies immediately deep (dorsal) to the fascia. The M. geniohyoideus of the iguanid liz- ards Amblyrhynchus, Brachylophus, CJialaro- don, Conolophus, Ctenosatira, Cychira, Dip- sosaurus. Iguana, Opiums, and Sauromalus extends posteriorly from the ventromedial surface of the mandibular rami and divides into medial and lateral bundles. The medial bundle passes posteriorly to insert on the ventral surface of the first ceratobranchial. The lateral bundle inserts on the ventrolater al surface of the first ceratobranchial, lateral to the medial bundle. It lies ventral and later- al to the anterior part of the M. hypoglossus. Oelrich (1956), in describing the condition in Ctenosaura, states: The lateral group twists so that at its origin the ventral surface is medial and the dorsal surface is lateral, the most lateral fibers extending dorsally and inserting later- ally. The more medial fibers fan out and insert all along the ventrolateral surfaces of the tongue to its posterior end, interdigitating with the dorsal transverse fibers of the intrinsic tongue musculature. The M. geniohyoideus of snakes is long and slender with one or more heads of origin. In Liotyphlops it arises as two heads, but in the Typhlopidae as a group its origin is from the inter-ramal connective tissue. In the Lepto- typhlopidae the origin is by a single head or tendon from the dentary. Rhinophis (Uropeli- tidae) has a medial head originating on the inter-ramal pad, but in Platyplecturus only the lateral head is present. Cylindrophis, San- zinia, Enhydris, Aipysusus, and Bothrops all possess a M. geniohyoideus with a single head. In most cases the M. geniohyoideus is bound to the tongue by a sheath and extends with the tongue at least to its base. In some forms such as Liasis, Eryx, and Xenopeltis the fibers extend even farther to insert on the M. hyoglossus. In Atretiwn the M. geniohyoideus resem- bles that of Varanus, with three divisions: lat- eral, ventral, and dorsal. Each of these origi- nates on the inter-ramal pad. The lateral division has two bundles, one of which ex- tends dorsolaterally to interdigitate with fi- bers of the second bundle. Together these bundles insert on the lining of the buccal floor. The ventral division extends post- erolaterally to separate into medial, inner, and a lateral bundle in the area of the glottis. The lateral group of fibers cross the medial ventrally to pass medially and to unite with the dorsal division of the M. geniohyoideus. The medial fibers extend posteriorly along the trachea to fan out and insert on the buc cal floor, with the main bundle inserting on the trachea itself. The fibers comprising the dorsal division of the M. geniohyoideus ex- tend posteriorly to insert on the lining of the buccal floor. The remainder of the muscle ex- tends posteriorly to join with the lateral bundle of the ventral division and pass paral- lel to the M. hyoglossus and insert into the tongue as a tendon. The M. geniohyoideus of Natrix arises from the inter-ramal ligament and consists of the M. geniohyoidei portiones minor and major. The portio major consists of fibers similar in configuration to the lateral bundle of the ventral division of Atretium. The medial bundle is not connected to the hyoid and may be a separate muscle, the M. mandibulotra- chealis as described in Natrix by Sondhi (1958). The portio major is similar to the dor- sal division oi Atretium, although its fibers do not insert on the buccal floor. The short, slen- der portio minor extends posteriorly to insert 334 Great Basin Naturalist Vol. 42, No. 3 on the anterior buccal floor. Its origin is adja- cent to that of the portio major. The portio major is long, and its fibers converge posteri- orly to insert on the base of the tongue. 2. Intrinsic Musculature The anatomy of the tongue is poorly un- derstood except in a few types that have been studied in detail. Attempts at explaining its morphology in Ctenosaura by Oelrich (1956) and Varanus and Matrix by Sondhi (1958) have only indicated the complexity of this structure in reptiles. The simplest inter- pretation is that the tongue of reptiles con- sists of a single muscle, the M. hyoglossus, which has been modified to serve many com- plex functions. In the Crocodilia the tongue lacks the rec- ognizable complex of intrinsic muscles seen in many and is formed from a more sim- plified association of the fibers of the hyog- lossus, which originates on the second cerato- branchials and inserts on the buccal floor. The tongue of Alligator has medial fibers of the M. hyoglossus that cross to opposite sides and interdigitate with fibers opposite the muscle. In Crocodylus the M. hyoglossus has a triple origin with fibers from the outer pro- ximal part of the second ceratobranchial, the ventral area of the second ceratobranchial at its point of articulation with the basihyoid, and the tendinous sheet where sternohyoid fi- bers insert on the articulation of the second ceratobranchial with the basihyoid. The tongue of Gavialis is described by Sondhi (1958) as having a M. hyoglossus with a double origin. One head originates as a ten- don from the middle of the ventral border of the second ceratobranchial, and the second head originates near the point of articulation between the second ceratobranchial and the basihyoid. The M. hyoglossus extends ante- romedially with interdigitations of fibers from both sides as the muscle inserts on the lining of the buccal floor. The tongue of Lissemys is formed by a M. hyoglossus consisting of two bundles each originating on the proximal portion of the second ceratobranchials (Gnanamuthu 1937). One bundle inserts on the side of the lingual process, and the other extends anteriorly to divide into two bundles to form the body of the tongue. In Trionyx the M. hyoglossus differs from that of Lissemys in that it is a single muscle as in Varanus and Matrix. The origin is from the ventral surface of the pro- ximal part of the second ceratobranchial in the form of longitudinal fibers. These extend anteriorly and are surrounded by a sheath of connective tissues. As the muscle passes ante riorly, the fibers split into three longitudinal bundles: outer, middle, and internal. This di- vision occurs anterior to the union of the two basal branches of the tongue. Sondhi (1958) describes the tongue of Va- ranus, using a series of successive transverse sections. To summarize his description, the longitudinal fibers of the M. hyoglossus be- come oblique and then transverse, with more and more longitudinal fibers changing direc- tion at the periphery of the tongue. The main muscular mass differentiates into two sets of fibers: one peripheral with circular fibers (pars externa) and one inner with longitudinal fibers (pars interna). The two groups are sep- arated by a thin fascial capsule. The circular fibers of the pars externa be- come tangential and interweave before the basal branches of the tongue combine at their dorsal borders. At the same time, the fibers in different areas of the pars externa change di- rections to form three intrinsic muscles: the Mm. verticalis linguae, transversalis linguae, and longitudinalis linguae. The M. verticalis linguae is composed of circular fibers of the pars externa on the inner side of each basal branch of the tongue, which extend vertically to lie between the remaining bundles of the pars externa. The dorsally dispersed fibers of the right and left pars externae become con- tinuous at the union of the two basal branches of the tongue to form the M. trans- versalis linguae. Posterior to the union of the two basal branches the M. longitudinalis ling- uae is formed from fibers of the M. trans- versalis linguae, along the dorsal branch of each half of the tongue, which change their direction from circular to longitudinal. Sever- al bundles of these fibers merge together to form a mass on the dorsolateral side of the tongue, which extends anteriorly to the apex. Just posterior to the anterior bifurcation of the body of the tongue, the pars interna of the M. hyoglossus on each side bifurcates to form two portions, which are separated by September 1982 Tanner, Avery: Buccal Floor of Reptiles 335 some of the bundles and the M. verticalis linguae. Each prong terminates with the dimin- ishing of the longitudinal and circular bun- dles and the insertion of their obliquely di- rected fibers on the epithelium of the tongue. Natrix (Xenochrophis) has been described by Sondhi (1958) as having a M. hyoglossus similar to that of the lizard Varanus. In Nat- rix the M. hyoglossus envelopes the second ceratobranchial at each side of the origin. Unlike that in Varanus, the M. hyoglossus of Natrix becomes ventromedial to the cerato- branchial and combines with its opposite member far posterior to the basihyoid. The M. hyoglossus also divides into parts externa and interna, but in the substance of the tongue rather than at its base as in Varanus. A number of longitudinal fibers of the M. hyoglossus separate from the rest of the pars externa at the periphery of the tongue to form the partes externa and interna. This change in direction of the fibers is directly associated with the formation of the Mm. verticalis linguae, transversalis linguae, and longitudinalis linguae the same as in Va ranus. The only difference is that the fibers of the M. longitudinalis linguae are formed more anteriorly in the body of the tongue in Natrix than in Varanus. Varkey (1979) describes the tongue of Ne- rodia cyckrprion as being formed of intrinsic tongue muscles and the M. hyoglossus. He describes the M. hyoglossus as long, slender, paired retractor muscles making up the bulk of the tongue. They arise from the medial edge of the posterior tips of the ceratobran- chials of the hyoid, pass rostrally, laterally, and ventrally to the intrinsic tongue muscles, and are pressed so closely together with them as to be almost indistinguishable. The hyog- lossus muscles attach to the hyoid comua, the tongue sheath, the oral mucosa, the fascia medial, and just posterior to the lateral sub- lingual glands. The intrinsic musculature of the tongue of Liduinura roseofusca has been described by Hershkowitz (1941) as consisting of five dis- tinct bimdles. In the posterior part of the tongue all but the M. verticalis are present. The M. transversus inferioris forms a sheet on the ventral side of the tongue extending dor- sally along the lateral side to meet the ven- tral extension of the M. transversus superioris. The M. transversus superioris occupies most of the dorsal part of the tongue deep to the superficial muscle, the M. lingualis, which is restricted to the most dorsal muscu- lar layer of the free, unforked part of the tongue. The M. verticalis forms a midsaggital ling- ual septum, thin toward the anterior and thick at the posterior end. The fibers of the M. verticalis run at right angles to those of the Mm. t. superioris and t. inferioris. Dor- sally its bundles interweave with those of the M. lingualis. The ceratoglossus muscles form a pair of central muscles extending the entire length of the organ and forming most of the cross sec- tion of the tongue. Posterior to the bifurcation of the tongue into terminal prongs, the Mm. verticalis ling- uae and transversalis linguae intersect at right angles. Thus, in section the tongue can be divided into four quarters composed of bundles of the M. longitudinalis linguae and the pars interna. At the anterior tip of the tongue, a dorsal and a ventral notch occur medially. The dor- sal notch deepens to separate the bases of the terminal prongs. At this point the bundles of the M. longitudinalis linguae of each side di- vide into smaller bundles and intermingle an- teriorly toward the tips of the prongs to ter- minate in the connective tissues of the lingual epithelium (Fig. 26). The M. hyoglossus in Chamaeleo brevi- cornis originates on the tip of the distal end of the first ceratobranchial. A small cartila- ginous knob on the end of the ceratobran- chial, which appears to be a remnant of the epibranchial, also serves as a point of origin for many fibers. The first and second comua extend anterolaterally from the basihyal; therefore the M. hyoglossus, in its contracted position, extends from its origin medially to the lingual process, where it makes a right angle turn to follow the lingual process into the tongue and to its insertion in the con- nective tissue surrounding the tongue. Upon reaching the tongue, the M. hyoglossus di- vides into the two sections described by Son- dhi (1958) as the pars externa and a medial longitudinal part, the pars interna. A series of circular fibers, which are a part of the sheath. 336 Great Basin Naturalist Vol. 42, No. 3 Fig. 28. Lateral view of the tongue of Chamaelon brevicarnis showing M. hypoglossus from its origin on the poste- rior comu to its folds before entering the tongue (BYU 12422). surroLuids the basal part of the tongue as a transverse sheet and encloses the distal ling- ual process and inserts dorsally into the tongue. In Chamaeleo the M. hyoglossus is folded, less so from its origin to the angle formed at its median posterior than as it extends along the lingual process (Fig. 28). The folds are deep and number 10 before the muscle enters the tongue. When fully extended, this folded part becomes an elongate, slender shaft sup- porting the clublike tongue. Gnanamuthu (1930) described the anatomy and function of the hyoid apparatus and tongue in Cha- maeleo cacaratus. His figures 5 and 6 corre- spond closely to our findings in Chamaeleo brevicomis. The folding is similar to the folds in the bellows of an accordian, whereas the muscular folds in free-tongued plethodontid salamanders is a series of looped folds (Tan- ner 1952). The outer bundle further divides into five to six smaller bundles, which lie beneath the dorsolateral border of the tongue to form the M. longitudinalis linguae. The fibers of the upper dorsolateral bundles of the M. longitu- dinalis linguae extend anteriorly to become obliquely transverse and give rise to the M. transversalis linguae, with the lower bundles continuing longitudinally to merge with each other. The internal longitudinal fibers of the M. hyoglossus become compact and vertical to form the M. verticalis linguae, just behind the tip of the tongue. At that point the middle bundle, between the Mm. trans- versalis linguae and verticalis linguae, passes dorsally so as to lie above the latter two bands. In the terminal end of each muscular September 1982 Tanner, Avery: Buccal Floor of Reptiles 337 CBl Fig. 29. Ventral view of hyoid apparatus and associated structures: A, Crotalus v. lutosus; B, Pituophis m. desert- icola. HG— m. hyoglossus, t— tongue. prong extending into the tongue from each In Figure 29 the general structural rela- side, the various bundles dwindle and insert tionships of the hyoid, tongue and M. hyo- in the subepithelial connective tissue of the glossus are depicted for the genera Crotalus tongue. «ind Pituophis. 338 Great Basin Naturalist Vol. 42, No. 3 In summary, the intrinsic muscles of the tongue are actually fibers of the hyoglossal muscles that extend in varying directions. Unfortunately, the remainder of our knowl- edge of the tongue and related structures is incomplete. Many structures such as lingual glands, glottis, trachea, and their associated muscles and nerves have not been fully in- vestigated in all groups. Zug (1971) has stud- ied the arterial patterns in iguanids, Winokur (1974) has studied the buccal mucosae in tur- tles, and Schumacher (1973) has examined the hyolaryngeal muscles and skeleton in tur- tles and crocodilians. VIII. Innervation of Buccal Floor Musculature A. General The innervation of muscles in reptiles has been generally neglected, and for this reason it is difficult to homologize their detailed musculature. Detailed descriptions of the nerve patterns in the buccal floor of reptiles are available from the following workers: Os- awa (1898), Watkinson (1906), Reese (1915), Willard (1915), Poglayen-Neuwall (1953, 1954), Oelrich (1956), Schumacher (1956, 1973), Sondhi (1958), and Rieppel (1978, 1981). Soliman (1964) describes and figures the nerves in the head of Chelydra serpentina and provides colored plates depicting the nerves entering the muscles associated with the buccal floor and the tongue. Islam (1955) and Islam and Ashig (1972) describe the cra- nium and cranial nerves of Uromastyx hard- wicki, and Renous-Lecru (1972) discusses the branchial plexus in Agama and Chalarodon. All these workers indicate that in reptiles the IXth (glossopharyngeal), Xth (vagus), Xlth (spinal accessory), and Xllth (hyoglossal) cranial nerves usually occur in close associ- ation and form a glossohyoidean plexus. Some uniformity does exist in the innervation of the throat muscles of reptiles, as demon- strated by the fact that in all reptiles the Vth (trigeminal) cranial nerve innervates the M. mylohoideus anterior, the Vllth (facial) in- nervates the Mm. mylohyoideus posterior and constrictor colli, and the Xllth (hypog- lossal) and anterior spinal nerves innervate the M. constrictor colli. B. Cranial Nerves Oelrich (1956) presented a clear picture of the pattern of cranial nerves in the iguanid Ctenosaura. He found the following nerves innervating muscles of the buccal floor. A similar pattern in all cases has been described for Anolis by Willard (1915), for the trige- minal in turtles by Poglayen-Neuwell (1953), and for Varanus by Watkinson (1906). 1. N. trigeminus: A branch of the trige- minal nerve (ramus ad musculum mylohyoi- deum) passes through the posterior mylo- hyoid foramen to enter the lateral fibers of the first mandibulohyoid muscle and termi- nates anteriorly on the M. intermandibularis posterior. A second branch, the anterior mylohyoid nerve, emerges on the medial side of the mandible from the anterior mylohyoid foramen of the splenial bone to pass over the M. mandibulohyoideus I to enter the ventral surface of the M. intermandibularis anterior. A section of the mandibular ramus continues anteriorly to the lingual ramus of the hypog- lossal nerve, where the latter passes through the anterior inferior alveolus foramen of the dentary to divide into two branches. The an- terior glandular branch passes the ventral surface of the M. intermandibularis anterior, whereas the posterior branch enters the Mm. intermandibularis anterior and genioglossus. 2. N. facialis: The facial nerve divides into a hyoid ramus that innervates a part of the M. intermandibularis that inserts on the ret- roarticular process of the mandible. It also in- nervates the M. constrictor colli and the pos- terior border of the M. intermandibularis. 3. N. glossopharyngealis: The M. hyog- lossus is innervated by a ramus formed from branches of the glossopharyngeal and hypog- lossal nerves. 4. N. hypoglossalis: There are four small ventral branches of the hypoglossal nerve that innervate the M. mandibulohyoideus I. The hypoglossal divides into three main branches at the point where the Mm. gen- ioglossus and hyoglossus join. These branches include the ramus lingualis lateralis, which extends anterolaterally to enter the insertion of the M. genioglossus and medial and lateral areas of the M. genioglossus. It next emerges to join the lingual ramus of the trigeminal September 1982 Tanner, Avery: Buccal Floor of Reptiles 339 nerve, which then enters the tongue. The re- maining two branches, intermediahs and me- diahs, go directly to the tongue, where they innervate its musculature. Watkins6n (1906) described the nerve pat- terns seen in Varanus and found the following: a. N. trigeminus: There are three main branches of the trigeminal nerve (rami op- thalmicus, maxillaris, and mandibularis); however, only the ramus mandibularis goes to the buccal floor, where it has three branches. The first branch, the ramus ad musculum mylohyoideus, originates from that part of the ramus mandibularis (portio alveolaris in- ferior) that lies within the alveolar surface of the dentary. It emerges to proceed posteri- orly, with branches going to the Mm. mylo- hyoidei posterior as profundus and superficialis. The second branch, the ramus muscularis et glandularis, also arises from the portio al- veolaris inferior of the mandibular ramus. A branch extends to the Mm. mentalis super- ficialis, mentalis profundus anterior, and mentalis profimdus posterior. It also enters the portiones major and minor of the M. genioglossus. A third branch, the ramus lingualis, origi- nates from the ramus mandibularis before the latter enters the alveolar canal. This branch emerges from the canal to pass along the ventral buccal floor, where it joins the ramus lingualis anterior of the hypoglossal nerve. It enters the lingual sheath and then the tongue, extending to the anteriormost extremity of the terminal prongs to innervate, with the hypoglossal nerve, the bundles of the M. hyoglossus. 2. N. facialis: The facial nerve emerges from the cranium and divides into an anterior branch, the ramus palatinus, and a posterior branch, the ramus hyomandibularis. Tlie lat- ter branch extends posteriorly as the ramus hyoideus to innervate the Mm. geniolateralis and constrictor colli. 3. N. hypoglossus: The hypoglossal nerve extends obliquely posterior along the dorsal side of the neck to the buccal floor, where it divides into two branches, each of which fur- ther subdivide into two branches. One branch forms the rami ad musculum geniotrachealis, and the second branch gives rise to the rami linguales anterior and posterior. The ramus ad musculum geniohyoideum extends obliquely over the M. ceratohyoideus to form two branches that innervate the mid- dorsal region of the M. geniohyoideus and lateral surface of the M. constrictor colli, respectively. The ramus ad musculum ceratohyoideum et musculum mandibulotrachealis extends from the M. interportalis to the M. cerato- hyoideus, innervating these and also sending branches to the Mm. cornuohyoideus and mandibulotrachealis. The third branch (ramus lingualis anterior) originates from the hypoglossal nerve and ex- tends along the lateral border of the tongue to eventually anastomose with the ramus lingualis of the mandibular ramus of the trigeminal nerve. As it does so, it sends branches to the sublingual glands and termi- nates in the M. genioglossus. A small branch also extends to both the Mm. gen- ioceratoideus and mandibuloproximalis. The ramus ad musculum mandibulohyoi- deum is derived from the hypoglossal nerve before the branching of the ramus lingualis posterior. It innervates the M. mandibulohyoideus. Two other branches derived from the hy- poglossal (lingual accessorii) innervate the posterior part of the base of the tongue. A fi- nal branch, the ramus lingualis posterior, is the terminal portion of the hypoglossal nerve. It also innervates the basal area of the tongue. Some information is available for other liz- ards such as Chamaeleo and Calotes (Gnana- muthu 1937), in which the formation of the lingual nerve varies. The lingual branch of the hypoglossal in Chamaeleo is separated from the glossohyoidean plexus and forms two branches, the rami linguale lateralis and medialis. The ramus lingualis lateralis extends posteriorly to innervate the M. genioglossus, and the main branch anastomoses with the lingual branch of the Vth cranial nerve; to- gether they penetrate the M. hyoglossus and join the ramus lingualis medialis that enters and innervates the M. hyoglossus. This same branch in the anterior region of the buccal floor unites with the combined lingual branch and with it also enters the tongue. In 340 Great Basin Naturalist Vol. 42, No. 3 Calotes the lingual branch of the hyoglossal nerve extends one branch to the M. gen- ioglossus and one to the tongue. The main branch unites with a ramus of the trigeminal to penetrate the tongue and there subdivides into many branches for innervation of the tongue muscles. Sondhi (1958) gives the following nerve pattern for the buccal floor of Matrix (Xe- nochrophis), a natricid snake: 1. N. trigeminus: The ramus mandibularis of the trigeminal nerve sends three branches to the buccal floor. The first branch (ramus ad musculum mylohyoideum) originates from the mandibular ramus immediately after the latter enters the alveolar canal of the dentary as the portio alveolaris inferior. It divides into two branches, one innervating the M. mylohyoideus posterior profundus and the other the M. mylohyoideus posterior. A second branch, the ramus muscularis et glandularis, originates from the portio alveol- aris inferior of the mandibular ramus. After emerging from the mandible, it extends me- dially to provide branches for the Mm. inter- maxillaris, genioglossus portio major, men- talis profundus anterior, and mentalis profimdus posterior. The third branch (ramus lingualis) arises from the portio alveolaris inferior of the mandibular ramus after the mandibularis et glandularis. It unites with the ramus lingualis of the hyoglossal nerve and extends medially to the lingual sheath and M. hyoglossus. 2. N. facialis: The facial nerve emerges from the foramen prooticum and extends to the M. mylohyoideus posterior as the ramus hyomandibularis, which has two branches to that muscle. 3. N. hypoglossal: The hypoglossal nerve has three main branches, including the ramus descendens that originates as a thin branch extending posteriomedially to innervate the ventral surface of the Mm. omohyoideus, sternohyoideus, and sternothyroideus. The second branch is the main stem of the hypoglossal nerve, which forms the ramus lingualis posterior. It extends forward as two branches, one entering the body and the other the base of the tongue. The third branch, the ramus ad musculum geniolateralis, originates in the hypoglossal nerve almost opposite the ramus lingualis posterior and innervates the M. gen- iolateralis. Distally the hypoglossal bifurcates into two branches, an inner ramus ad muscu- lum mandibulotrachealis and an outer ramus ad musculum geniohyoideum. The inner divi- sion extends anteriorly and medially to in- nervate the posterior part of the M. man- dibulotrachealis. The outer branch extends anterolaterally to form two branches that in- nervate the M. geniohyoideus. Langebartel (1968) has summarized the in- nervation of the muscles of the buccal floor in other snakes. The mandibular division of the trigeminal nerve innervates the M. inter- mandibularis and parts of the tongue. The fa- cial innervates part of the Mm. constrictor colli and the cervicomandibularis and sends some branches to the tongue. Some branches from glossopharyngeal and the vagus in- nervate the M. ceratomandibularis, but only one branch innervates the M. hyotrachealis. The hypoglossal nerve innervates the Mm. geniohyoideus, ceratomandibularis, and ster- nohyoideus. The Mm. genioglossus and hy- poglossus are innervated by an anterior branch of the hypoglossal nerve. Very com- monly the glossopharyngeal, vagus, and hy- poglossal nerves combine to innervate the lingual sheath and the Mm. genioglossus and hypoglossus. The hypoglossal may also have anterior and posterior branches that enter the tongue. Last, an anterior branch of the hypo- glossal unites with a branch of the trigeminal to innervate the Mm. genioglossus and gen- iotrachealis. Varkey (1979) describes the in- nervation of muscles in Nerodia, but does not attempt to identify the nerves. Soliman (1964) describes and figures the cranial nerves of Chelydra serpentina. Col- ored plates depict the various nerves entering the muscles associated with the buccal floor and the tongue. Trionyx has been described by Sondhi (1958), who indicates the existence of the fol- lowing nerve pattern, comparable to that re- ported for Chelydra: 1. N. trigeininus: The mandibular ramus of the trigeminal nerve has two branches, in- cluding the ramus ad musculum mylohyoi- deum that arises in Varantis and Matrix, and the ramus lingualis. The former branch ex- tends posteriorly along the medial side of the mandible to divide into two branches, one of September 1982 Tanner, Avery: Buccal Floor of Reptiles 341 which innervates the M. mylohyoideus ante- rior and the other which innervates the M. mylohyoideus posterior. The ramus hnguaUs arises from the portio alveolaris inferior and emerges from the mandible through a small foramen to innervate the M. genioglossus. Sohman and Hegazy (1971) also describe this nerve in Chalcides ocellatus. 2. N. facialis: The facial gives rise to the ramus hyomandibularis, which innervates the buccal floor. It extends posteriorly as the ramus digastricus and sends a branch to the M. constrictor colli and another to the M. constrictor superficialis. 3. iV. hypoglossus: The hypoglossal nerve extends along the anterior part of the neck to the M. ceratohyoideus, where it gives rise to two branches, the rami descendens and ad muscukun stemothyroideum. A third branch (ramus ad musculum geniohyoideum) is formed as it emerges on the ventral side of the M. ceratohyoideus. Finally, it extends an- teriorly to provide the ramus lingualis and then terminates by dividing into two branches, the rami ad musculum entoglos- sohypoglossalis and ad musculum hy- poglossolateralis. The ramus descendens extends anterome- dially beyond the second ceratobranchial to form two branches that innervate, respective- ly, the Mm. omohyoideus and the ster- nohyoideus. The ramus ad musculum ster- nothryoideum extends across the surface of the Mm. omohyoideus and sternohyoidevis to innervate the M. sternothyroideus. The ramus ad musculum ceratohyoideus extends to the dorsal surface of the M. ceratohyoideus, which it innervates. The ramus ad musculum mandibulohyoideum is a small branch ex- tending anteriorly to innervate the ventral surface of tlie M. mandibulohyoideus. The ramus ad musculum geniohyoideum extends anteriorly to form two branches, with one in- nervating the portio ventralis and the other entering the portio distalis of the M. gen- iohyoideus. The ramus lingualis extends me- dially to enter the base of the tongue, where it passes anteriorly inside the tongue to in- nervate the M. hyoglossus. As in Lissemys, there are no anastomoses with the lingual branch of the trigeminal. The ramus ad mus- culum entoglossohypoglossalis is a delicate branch innervating the M. entoglossohypo- glossalis. Finally, the ramus ad musculum hy- poglossolateralis extends obliquely lateral to innervate the M. hypoglossolateralis. Sondhi (1958) has investigated the nerve patterns of the buccal floor seen in Gavialis and presents the following pattern. 1. N. trigeminus: The mandibular ramus of the trigeminal nerve forms two branches, the rami ad musculum mylohyoideum and lin- gualis. The former emerges from the dentary and passes posteriorly to innervate the dorsal surface of the M. constrictor colli. The ramus lingualis emerges from a foramen after aris- ing from the portio alveolaris inferior. It passes obliquely anterior to innervate the M. genioglossus portio major. 2. N. facialis: The ramus hyomandibularis of the facial nerve sends a branch, the ramus hyoideus digastricus, of Sondhi, posterior to the neck to divide into two branches. The first branch innervates the M. constrictor pharyngis and the second extends dorsally to the Mm. constrictor colli and constrictor superficialis. 3. N. hypoglossus: On the dorsal side of the neck the hypoglossal nerve divides into four branches. The first branch, or ramus de- scendens, divides into two branches at or near the middle of the M. omohyoideus. These branches innervate the M. omo- hyoideus and the M. sternohyoideus, respec- tively. The second branch, ramus ad muscu- lum sternohyoideum, passes obliquely posterior to divide into several branches that innervate the M. sternohyoideus. The ramus lingualis posterior forms the third branch and sends a subdivision, the ramus ad musculum geniohyoideum, to the M. geniohyoideus, and other branches enter the tongue and in- nervate the M. geniohyoideus portio major. The last branch, ramus lingualis anterior, ex- tends posteriorly to the mandible to in- nervate the Mm. ceratohyoideus and mandi- bulohyoideus, and other branches extend anteriorly to enter the tongue and the Mm. genioglossi portiones minor and major. Reese (1915) indicates that the ramus man- dibularis (ramus maxillaris inferior of Reese) of the crocodile divides into two and then four branches. Two of these branches in ner- vate the M. mylohyoideus. The M. hyoglossus is served bv branches of the IXth and Xllth 342 Great Basin Naturalist Vol. 42, No. 3 nerves. The hypoglossal nerve also sends branches to the Mm. omohyoideus, ster- nohyoideus, geniohyoideus and ge-nioglossus. B. Spinal Nerves Oelrich (1956) reports that in Ctenosaura the first spinal nerve innervates the ventral part of the M. omohyoideus and the dorsal part of the stemohyoideus. Sondhi (1958) in- dicates that in Varanus and Matrix the united stems of the first and second spinal nerves anastomose with the hypoglossal nerve and extend posteriorly in the neck to send small branches to the Mm. stemohyoideus and ster- nothyroideus and a large branch to the M. omohyoideus. Some of the succeeding spinal nerves also innervate the M. constrictor colli. In Natrix, as in Varanus, the first and sec- ond spinal nerves innervate parts of the M. constrictor colli. In some other snakes many spinal nerves innervate the Mm. neuroman- dibularis, costomandibularis, costo-cutanei in- ferior and superior, omohyoideus, ster- nohyoideus, and transverus branchialis. In Trionyx, Gavialis, and Crocodylus the united stem of the first and second spinal nerves irmervates the M. constrictor colli, whereas in Crocodylus numerous branches of first, second, and third spinal nerves in- nervate the smaller ventral muscles of the neck. IX. Discussion An examination of the preceding descrip- tions show that the information on the hyoid and associated structures was widely scat- tered and incomplete. Although morphology is one of the oldest branches of biology, there is an absence of complete accounts of the gross anatomy of the buccal floor of reptiles as a class. Similar gaps in our knowledge exist for other anatomical areas of the reptilian body. In spite of our acceptance of some rep- tilian ancestral stocks as being the lines of de- scent for birds and mammals, anatomists have not vigorously pursued studies to show phy- logenetic relationships. The lack of a com- plete understanding of these groups is as- tounding considering the important phylogenetic position of reptiles. Despite the lack of information, some gen- eralizations can be made. As indicated by Sondhi (1958), the buccal floor in many rep- tiles has three functions: (1) it participates in the act of inspiration and expiration, (2) it aids in the capture and the deglutition of food, and (3) it provides the mechanisms of tongue movement. To Sondhi's list should be added two additional functions: (4) behav- ioral display and (5) sensory reception. The important role of the buccal area as a respiratory throat pump has been explored by Gnanamuthu (1937), who demonstrated the part played in Hemidactylus. He states. The contraction of the thorax expeUing air would result in the inflation of the buccal cavity, and when next the thorax relaxes this impure air may be taken into the lungs again, because the thoracic contraction and expan- sion follows each other so rapidly. However, the eleva- tion of the rnouth floor and tongue through the aid of transverse and hyoid muscles just when the thorax con- tracts serves to expel the vitiated air effectively out of the body. Respiratory mechanisms in reptiles vary widely. Calotes utilizes the limbs of the ante- rior cornua and the attached muscles to ac- tively raise and lower the throat. The posi- tions of cornua and ceratobranchials and associated muscles in Varanus indicate a change of the volume of the throat caused by dilation and compression of the floor of the mouth. Among the testudines, the posterior part of the M. hyoglossus and the entire Mm. cerato- hyoideus, entoglossohypoglossalis, and hypo- glossolateralis utilize the jointed basihyoid and hypoglossum to move the throat up and down as one solid piece. Although these structures may not be important in respira- tion (Mitchell and Morehouse 1863), there is reason to believe that both aquatic and ter- restrial turtles pump the throat to exchange water and air in the nasal canals and buccal cavity for sensory functions (McCutcheon 1943). In Figure 19 we attempt to reproduce the extensive fimbriations on the bucco- pharyngeal floor of Trionyx. The total func- tion of these numerous filaments may not be fully understood, but seemingly they are im- portant in aquatic respiration (Girgis 1961). In snakes, inspiration and expiration are accomplished by the muscles of the body wall compressing the lungs for expiration and September 1982 Tanner, Avery: Buccal Floor of FIeptiles 343 expanding for inspiration. A minor contribu- tion is made by the expansion and con- traction of the anterior part of the body. As a result, the hyoid has become greatly reduced and contributes mostly as a support for the buccal floor and as a support for the muscles and membranes that open and close the glottis. For further information on respiration in vertebrates, see Hughes (1963), Gans and Hughes (1967), Bishop and Foxon (1968), and Gans (1969). Food capture and deglutition in reptiles is difficult to correlate with the morphology of the buccal floor. For example, snakes have a ligamentous connection between the man- dibular rami and movable articulations of the maxilla, palatine, pterygoid, and quadrate, which allow for the movement of one side of the jaw apparatus to move forward and se- cure a firm hold on the prey before moving the other side, as indicated by Gans (1961) and Frazzetta (1966). Such a situation does not exist in the Lacertilia, Amphisbaenia, Rhychocephalia, Testudines, or Crocodylia, making comparisons difficult, if not impos- sible. In the latter three groups, however, the food capturing and swallowing mechanisms are basically similar owing to the greater similarity of throat anatomy. The movement of the tongue is important in most reptiles because of its sensory nature and association with Jacobson's organ. The tongue is simplest in the primitive testudines and Crocodilia, indicating a more ancient and conservative nature in these groups. The primitive lizards, such as iguanids, and some testudines, such as Gopherus, have a thick, fleshy tongue, used both for sensory activities and manipulation of the food within the mouth (Avery and Tanner, 1971; Gnana- muthu, 1937). An advanced lizards, such as Varanus, the tongue is similar to that of snakes in gross morphology. The fact that the associated throat musculature in these two groups differs is an indication that perhaps the manipulation of the tongue in varanids and snakes has been, at least partially if not completely, freed from the buccal floor musculature. Last, the buccal floor has behavioral impli- cations in many lizards, particularly the igua- nids, in which males often have enlarged throat dewlaps. The behavioral implications of these structures is beyond the scope of this paper [see work of Carpenter 1965 (Anolis), 1967, 1977 (Iguanids), 1970 (Agamids)], but in the forms with the best developed dew- laps, such as Anolis and Iguana, the second ceratobranchials and associated musculature provide the main structural components of movement. Some generalizations about the buccal floor can also be made. The more primitive the organism, the less complicated and spe- cialized the gross anatomy of the buccal floor. This is apparently true for most orders, although there are exceptions within some orders (such as in some testudines). In the primitive forms, the hyoid has retained more cornua, some specialized muscles are absent, and the tongue is less differentiated. In the more advanced forms, such as lizards, the hyoid has become complex and the muscula- ture has proliferated and specialized. Lizards show a greater variation in the morphology and function of the tongue than do other groups of reptiles. Tongues are structured for food manipulation (Iguanidae and Amphisbaenia), food getting (Cha- maeleonidae and Amphisbaenia), and also for sensory functions in such groups as Cnemido- phorus, Heloderma, and Varanus. Such func- tional variations have in turn altered the bas- ic morphology of the buccal floor to accommodate the adaptive feeding habits and the associated sensory and behavioral ac- tivities. In snakes specializations of feeding and life habits have caused a secondary re- duction of many elements of the buccal floor, particularly in the skeletal structures, and the tongue is no longer a manipulator of food. In snakes the tongue is filamentous and impor- tant primarily as a sense organ. As indicated by Sondhi (1958), there is a structural sim- ilarity between the tongues of some lizards (Varanus) and snakes. This, Sondhi reasoned, may have led to the development of the highly sensitive tongues of snakes. At least, such lizards have a flexible tongue and the terminal forking is structurally similar enough to suggest an evolutionary relation- ship. Perhaps this is an example of con- vergence of structure to perform a similar function in distantly related groups. In general it is difficult to draw major phy- logenetic conclusions from the buccal floor 344 Great Basin Naturalist Vol. 42, No. 3 because the scope of such a study is neces- sarily hmited to one specialized area and can be misleading. When hyoid elements are lost, the associated muscles are also lost or may become unrecognizable. Thus the implication of presence or absence of structures is also misleading. Future morphological phyloge- netic studies in the area of the buccal floor should be supplemented by embryological in- formation, as indicated by DeBeer (1930, 1951) and Edgeworth (1935). Such research will provide clues as to which structures have been lost, fused, readapted or never possessed by an organism. Acknowledgments We wish to thank those who have aided us during the preparation of this study, particu- larly our own institutions who have provided space and secretarial help and have permit- ted us to dissect materials from their collec- tions (Brigham Young University Life Science Museum, BYU, and Southern Connecticut State College, SCSC). We have also received study materials from the following: Dr. George R. Zug, U.S. National Museum; Dr. Carl Cans, University of Michigan; Dr. Alan E. Leviton, California Academy of Sciences; Dr. Richard G. Zweifel, American Museum of Natural History, and Mr. Jose P. Rosado, Museimi of Comparative Zoology. We re- ceived, on exchange, specimens from Dr. F. H. Rehmani, Zoologische Museum, Kanpur, India. These were very helpful as com- parative materials, and for their use we are indeed grateful. 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The distribution and patterns of the major arteries of the iguanids and comments on the intergeneric relationships of iguanids (Rep- tilia:Lacertilia). Smithsonian Contrib. Zool. 85:1-23. WESTERN DIAMONDBACK RATTLESNAKE IN SOUTHERN NEVADA: A CORRECTION AND COMMENTS Frederick H. Emmerson' Abstract.— Reports of the occurrence of Crotahis atrox Baird & Girard from Nevada are reviewed. There is no evidence to support the occurrence of this species in Nevada. Andersen and Emmerson (1970, Great Ba- sin Nat. 30:107) reported the taking of a single specimen of western diamondback rat- tlesnake, Crotahis atrox Baird & Girard, from west of the Colorado River in Clark County, Nevada. The snout-vent length was erroneously re- ported as "163 mm," permitting the infer- ence that a breeding population may have been present in the Searchlight area, result- ing in the collection of a neonate. In fact, the specimen (Univ. of Nevada, Las Vegas R5118) was an adult male with a snout-vent length of 1054 millimeters. During the years subsequent to the above- cited report, and as recently as 1979, a num- ber of representations were made to me that "diamondbacks" had been taken or seen in Clark County. Inasmuch as no one making the claims produced a specimen of C. atrox, nor was any report offered by a competent observer, I conckided that the reports were fictitious or were based upon sightings of C. mitchelli or scutulatus. I suggest, therefore, that caution be exercised in considering Cro- tahis atrox as part of the Clark County, Ne- vada, herpetofauna, at least until a series of specimens is at hand. 'Utica Zoo, Steele Hill Road, Utica, New York 13501. 350 PREVALENCE OF ELAEOPHORA SCHNEIDERI AND ONCHOCERCA CERVIPEDIS IN MULE DEER FROM CENTRAL UTAH Lauritz A. Jensen', Jordan C. Pederson-, and Ferron L. Andersen' .\bstr.\ct.— Thirteen of 265 deer (4.9 percent) from central Utah were positive for Elaeophora schneideri, and 180 (67.9 percent) were infected with Onchocerca cervipedis. The rate of infection for E. schneideri and O. cervipedis in- creased significantly with age of the host (chi-square of 17.5 and 15.5, respectively, p < 0.005). The lack of elaeopho- rosis in elk from the region is presumably due to the low density of the parasite in mule deer. The arterial worm (Elaeophora schneideri) is enzootic in mule deer [Odocoileus he- tnionus), elk {Cervus canadensis), and other ruminants in the Rocky Mountain area (Hib- ler and Adcock 1971, Worley 1975), although the prevalence and geographic distribution in Utah is largely undetermined. From 1978 to 1981 biologists of the Utah Division of Wild- life Resources (DWR) identified the arterial worm in one calf and one bull elk from southern Utah and detected microfilariae of E. schneideri in 19 of 37 mule deer from the same locality (Coles 1982, pers. comm.). Ad- ditionally, from 1978 to 1981 DWR per- sonnel recovered the filarial nematode in three blind moose and one moose with nor- mal sight in northern Utah (Babcock 1982, pers. comm., Kimball 1982, pers. comm.). In contrast, of 23 adult elk and seven calves ex- amined by the authors and DWR officers in central Utah during summer 1981, none manifested clinical signs of elaeophorosis (un- publ. data). Mule deer are apparently the normal de- finitive host for E. schneideri, and the adult worms are predominately located in the ar- teries of the neck. Elk and moose, on the other hand, represent abnormal hosts, and are severely affected by the larval stages found in the cephalic arteries, arterioles, and capil- laries. Heavily infected elk calves frequently die 7 to 10 days after infection (Hibler 1981). Other symptoms of elaeophorosis in elk and moose include damage to the central nervous system, nystagmus, blindness, cropping of the ears, deformity of the antlers, necrosis of the muzzle and nostrils, and emaciation (Hibler and Adcock 1971, Worley et al. 1972). The legworm (Onchocerca cervipedis) is another filarial nematode enzootic in mule deer in the Rocky Mountains (Walker and Becklund 1970). Adults of the legworm may cause inflammation in the subcutaneous tis- sue (Wehr and Dikmans 1935); however, in- fections generally are not clinically signifi- cant (Senger 1963). The present study was undertaken to determine the occurrence of E. schneideri and O. cervipedis in deer from central Utah. Samples of skin from the muzzle of 265 deer were macerated in physiological saline and examined for microfilariae of E. schneid- eri and O. cervipedis following generally the procedure described by Weinmann et al. (1973). The animals were hunter-killed in au- tumn 1981 in the counties of Carbon, Duchesne, Emery, Juab, Sanpete, Utah, and Wasatch. Thirteen of the 265 deer (4.9 percent) were positive for E. schneideri and 180 (67.9 per- cent) were infected with O. cervipedis (Table 1). Deer positive for both parasites were re- covered from all counties listed above. The rate of infection for the arterial worm and the legworm increased significantly with age of the host (chi-square of 17.5 and 15.5, re- spectively, p < 0.005). The apparent lack of elaeophorosis in elk in central Utah may be related to the low density of the parasite in mule deer from the region. The high rate of infection for both species of roundworms demonstrated in older 'Department of Zoology, Brighatn Young University, Prove, Utah 84602. ■Utah Division of Wildlife Resources, 1115 North Main. Springville, Utah 84663. 351 352 Great Basin Naturalist Vol. 42, No. 3 Table 1. Prevalence of Elaeophora schneideri and Onchocerca cervipedis in mule deer from Central Utah. Deer infected with E. schneideri Number Age of of deer deer examined Number Percent Number Percent Deer infected with O. cervipedis Yearlings 175 2-3 years 27 >3 years 63 Total 265 2 1.1 105 60.0 2 7.4 21 77.8 9 14.3 54 85.7 13 4.9 180 67.9 animals is presumably related to the time of exposure in the enzootic area. Yearlings are apparently not refractory to either parasite. Voucher specimens: microfilariae of E. schneideri and O. cervipedis USNM Helm. Coll. No 76931. Acknowledgments Thanks are given to Holly Betteridge, Har- old Blackburn, Brad Bradly, Paul Tervort, and Dick Worthen, Division of Wildlife Re- sources, Utah, for assisting with the collecting of the samples. We are indebted to Dr. Charles P. Hibler, Wild Animal Disease Cen- ter, Colorado State University, for confirming the identifications of the microfilariae. Ap- preciation is also extended to Kimball T. Harper, Rodney T. John, and Bmce L. Welch for their suggestions in the preparation of this manuscript. Literature Cited Babcock, W. H. 1982. [Letter to Jordan C. Pederson]. Located at: Utah Division of Wildlife Resources, Springville, Utah. Coles, F. H. 1981. [Letter to Jordan C. Pederson]. Lo- cated at: Utah Division of Wildlife Resources, Springville, Utah. Hibler, C. P. 1981. Elaeophorosis. Pages 53-59 in W. J. Adrian, ed.. Manual of common wildlife diseases in Colorado. Colorado Division of Wildlife. Hibler, C. P., and J. L. Adcock. 1971. Elaeophorosis. Pages 263-278 in J. W. Davis and R. C. Ander- son, eds.. Parasitic diseases of wild mammals. Iowa State Univ. Press, Ames, Iowa. Kimball, J. F. 1982. [Letter to Lauritz A. Jensen]. Lo- cated at: Utah Division of Wildlife Resources, Springville, Utah. Senger, C. M. 1963. Some parasites of Montana deer. Montana Wildl. Autumn: 5-13. Walker, M. L., and W. W. Becklund. 1970. Checklist of the internal and external parasites of deer, Odocoileus hemionus and O. virginianus, in the United States and Canada. Special Publication No. 1, Index-Catalogue of Medical and Veter- inary Zoology, U.S. Govt. Print. Off., Washing- ton, D.C. Wehr, E. E., and G. Dikmans. 1935. New nematodes (Filariidae) from North American ruminants. Zool. Anz. 110:202-208. Weinmann, C. J., J. R. Anderson, W. M. Longhurst, and G. Connolly. 1973. Filarial worms of Co- lumbian black-tailed deer in California. 1. Obser- vations in the vertebrate host. J. Wildl. Dis. 9:21.3-220. Worley, D. E. 1975. Observations on epizootiology and distribution of Elaeophora schneideri in Montana ruminants. J. Wildl. Dis. 11:486-488. Worley, D. E., C. K. Anderson, and K. R. Greer. 1972. Elaeophorosis in moose from Montana. J. Wildl, Dis. 8:242-244. ECOMORPHOLOGY AND HABITAT UTILIZATION OF ECHINOCEREUS ENGELMANNII AND E. TRIGLOCHIDIATUS (CACTACEAE) IN SOUTHEASTERN CALIFORNIA Richard I. Yeaton' Abstract.— The relationship between form and habitat utihzation of Echinocereus engebnannii and E. triglochi- diatiis was studied in southeastern CaUfomia. The major difference in form is in the density of stems comprising the canopy. These differences in morphology create differences in the daily heat loads of each species. Echinocereus trig- lochidiatus, with its stems densely packed and in contact with each other over much of their lengths, continues to gain heat internally as the sun sets due to conductance between the stems. In contrast E. engelmannii, with a more open canopy, begins to lose heat as the sun goes down. As a result, E. triglochidiatus is successful in the juniper-pin- yon zone where winter temperatures are cold for long periods and E. engelmannii is more successful in the lower desert regions where very hot, summer temperatures prevail. In the latter case, daytime buildup in heat load is re- duced by convective cooling as air moves through the open canopy. Differences in microhabitat utilization occur that provide a second mechanism to reduce daily heat load buildup on hot summer days in the juniper-pinyon zone. Plants in their natural habitats must cope with the environmental variations expe- rienced in each microhabitat. A series of adaptive strategies, which involves gaining access to sufficient water, nutrients, and light for maintenance, growth and reproduction while at the same time avoiding the effects of desiccation and extreme temperatures that can lead to mortality, is used to achieve this (e.g.. Gates 1962). Nowhere are these strate- gies more apparent than in plants occupying extreme environments. In particular, most morphological aspects of members of the Cactaceae have been suggested as represent- ing ecological strategies for their successful adaptations to hot, dry environments (Gates 1962, Felger and Lowe 1967, Nobel 1978, 1980a, Yeaton et al. 1980). The literature on problems of adaptations in form for Cactaceae involves two types of studies; one in which access to sufficient light for photosynthesis is considered (Rodriquez et al. 1976, Nobel 1980b, Yeaton et al. in re- view) and the other in which avoidance of extreme temperatures, particularly freezing temperatures, is emphasized (Felger and Lowe 1967, Gibbs and Patten 1970, Mozingo and Comanor 1975, Nobel 1978, 1980a). Two extremes in the form of Cactaceae parallel this division. When access to light is dis- cussed, platyopuntias, a group with flattened cladodes, are usually studied. In contrast, when temperature is a problem, the growth form of the species studied is usually some variant of a cylinder. In this study, the relationship between heat load and habitat utilization is examined for two species of Echinocereus that are mor- phological variants of the cylindrical form. Echinocereus engebnannii (Parry) Lemaire is found on rocky slopes of elevations of 600-1500 m in the Mojave Desert of Califor- nia (Benson 1969). It is caespitose with 5 to 60 stems forming an open mound. The sec- ond species, Echinocereus triglochidiatus Engelm., is found at somewhat higher eleva- tions (1000-2500 m) above the deserts, usual- ly in the juniper-pinyon woodland (Benson 1969). It also has a caespitose form with mul- tiple stems forming a dense mound. The com- pactness of stems in the mounds is the most striking difference in form between the two species and results in a lower effective sur- face-to-volume ratio in E. triglochidiatus. I concentrated on this morphological dis- similarity (1) to determine what differences occur in the daily heat load experienced by individuals of each species and (2) to explain 'Centre de Botanica y Centro Regional para Estudios de Zonas Aridas y Semidridas, Colegio de Postgraduados, Chapingo, Edo. de Mexico, Mexico. Pres- ent address: Molecular Biology Institute, UCLA, Los Angeles, California 90024. 353 354 Great Basin Naturalist Vol. 42, No. 3 differences in habitat utilization by each species. Study Areas and Methods A population of Echinocereus engelmannii was studied at Hole-in-the-Wall at the base of the Providence Mountains in the Mojave Desert of southeastern California (latitude 35°3' N, longitude 115°23' W). For a de- scription of this site, see Yeaton and Cody (1979). Populations of E. engelmannii and E. triglochidiatus were studied in the juniper- pinyon zone (latitude 35°4' N, longitude 115°28' W). Here rainfall is more abundant and average maximum daily temperatures are lower (Costing 1956, Trombulak and Cody 1980). As a result, snow is common during the months of December, January, and February and persists for several days or weeks, in contrast to the lower desert site. At the upper site the dominant vegetation con- sists of Artemisia tridentata, Juniperus mon- osperma, and Pinus monophylla. At each study site, all individuals of E. engelmannii and E. triglochidiatus were cate- gorized according to the kind of microhabitat utilized. The characteristics of these micro- habitats were quantified from the perspective of the plant. Three general microhabitats may be distinguished at the jujiiper-pinyon site: rocky slope, composed of a mosaic of boulders and gravel and found at slope angles greater than 8 degrees; under juniper and pinyon, generally at slope angles less than 8 degrees; and washes, disturbed areas with Ar- temisia tridentata dominating the plant com- munity. At the desert study site, only the rocky slope microhabitat is utilized exten- sively, washes are rarely colonized by Ech- inocereus (possibly due to greater effects of erosion), and no counterpart to juniper- pinyon canopy exists. The following micro- Table 1. Habitat utilization by individuals of Echino- cereus triglochidiatus and E. engelmannii in the juniper- pinyon zone of the Providence Mountains, California. Pinyon pine Species Rocky slopes or wash E. triglochidiatus E. engelmannii 65 38 39 0 Fisher Exact Probability Test, p < 0.001. habitat characteristics were recorded for each Echinocereus individual whenever pos- sible: slope aspect (either north-facing [270-90 degrees] or south-facing [90-270 de- grees]), shaded or unshaded by adjacent rocks or plants. These data were organized into contingency tables with row headings con- sisting of the two species of Echinocereus and column headings describing the contrasting characteristics of each microhabitat. Entries into the tables were the numbers of individ- uals encountered in each situation. Totals from table to table were not necessarily equivalent because some individuals could not be assigned to a particular category. Ei- ther Chi-square or Fisher Exact Probability tests were employed to determine differences in microhabitat utilization between the two species (Siegel 1956). The sizes of the 10 largest individuals of each species in each microhabitat were mea- sured. Because both species form mounds that are roughly hemispherical in shape, one approximation of size is the diameter of the mound. A second measure of size is the num- ber of stems comprising each mound. To de- scribe the degree of openness of the canopy of each individual, a ratio of the number of stems divided by the mound diameter was calculated and compared by means of a Stu- dent's t-test (Steel and Torrie 1960). In addi- tion, the diameters of one stem from 25 indi- viduals of each species were measured and tested similarly. Daily temperature regimes were measured for a large (0.5 m diameter) individual of each species that had been transplanted into a shallow clay pot and removed to an open site where the plant would not be shaded over the course of a day. The two species were set side-by-side and a Yellowsprings In- strument Tele-thermometer and probes were used to simultaneously record hourly ambient air temperature, cactus surface temperatures (on the east-facing side of the center stem), and internal stem temperatures (at 6 cm and 10 cm depths in the center stem of each indi- vidual). These temperatures are plotted for 15 September 1980 and are used to illustrate the differences in daily heat loads due to de- gree of openness of the canopy. Graphs were made of the daily course of the ambient air September 1982 Yeaton: Cactus Habitat Utilization 355 temperatures and the differences between ambient temperatures and surface and inter- nal temperatures for each species. Results At die desert site, only individuals of Ech- inoceretis engelmannii, utilizing rocky slope microhabitats, are found. At the juniper- pinyon site, individuals of both species occur. At this site E. engelmannii is found only in the rocky slope microhabitat, but E. trigloehi- diatus is foimd in all tliree microhabitats (Table 1). At the juniper-pinyon site, E. eng- elmannii is located on south-facing slopes only, but individuals of E. triglochidiatits are equally divided between north- and south- facing slopes (Table 2). Echinocereus engel- mannii apparently requires open, sunny mi- crohabitats for successful establishment, but E. triglochidiatus is favored by more shaded, northern exposures. This distinction is further illustrated by the characteristics of micro- habitat involving shading. E. engelmannii is almost always found in unshaded micro- habitats, but E. triglochidiatus is found in shaded situations, whether it occurs on north- or south-facing slopes (Table 3). The means and standard errors for size measurements of tlie 10 largest individuals of each species in each microhabitat are given in Table 4. Comparison of the mean values for moimd diameters and number of stems between the two species on the rocky slope at the juniper-pinyon site shows that E. trig- lochidiatus is much larger than E. engel- mannii (for diameter t = 6.54, d.f. = 18, p < 0.001; for number of stems t = 10.23, d.f. = 18, p < 0.001). These differences combine to give significantly lower mean values for the ratio of stem number/ mound diameter for E. engelmannii, indicating that the spacing between stems is relatively large (t = 12.58, d.f. = 18, p < 0.001). This is not due tQ differences in stem diameter because no significant difference was found between the two species (for E. engelmannii stem di- ameter X ± S.E. = 5.18 cm ± 0.16; for £. triglochidiatus x ± S.E. = 4.82 cm ± 0.15; t = 1.63, d.f. = 48, 0.2 < p < 0.1). No differ- ence in the stem number/mound diameter ratio exists between the upper and lower populations of E. engelmannii; however indi- viduals of the upper population are signifi- cantly smaller in diameter (t = 6.06; d.f. = 18, p < 0.001) and in stem number (t = 6.38, d.f. = 18, p < 0.001) than are individ- uals measured at the desert site. For E. trig- lochidiatus a gradual reduction in diameter and stem number and an increase in the openness of the canopy occurs from the rocky slope through the juniper-pinyon to the washes (Table 4). The only significant dif- ferences occur between the rocky slope and wash microhabitats for this species (for mound diameter t = 2.47, p < 0.05; for stem number t = 4.48, p < 0.001; for the ra- tio t = 3.53, p < 0.01; d.f. = 18 in all cases). The daily temperature regimes are shown in Figure 1. Ambient air temperature in- creases from early morning until late after- noon and decreases as the sun sets (Fig. la). For the relatively open-canopied E. engel- m.annii, surface temperatures are much greater during the early part of the day and decrease rapidly as the east-facing side of the stem is shaded in the later daylight hours (Fig. lb). At the 6 cm depth, temperatures appear buffered and fluctuate around the ambient temperature during the day, heating up as the sun goes down (Fig. lb). At the 10 cm depth, temperatures start below ambient in the morning, increase rapidly during the day, and gradually decrease in the evening (Fig. lb). In contrast, the closed-canopied E. triglochidiatus maintains surface and internal stem temperatures below those of ambient during the daylight hours and gradually in- crease as the sun sets. By midnight, temper- atures in E. triglochidiatus stems are greater than those for E. engelmannii under the same conditions (Fig. Ic). Table 2. Slope aspect utilization for individual of Echinocereus triglochidiatus and E. engelmannii in the juniper-pinyon zone of the Providence Mountains, California. North-facing South-facing Species (270-90 degrees) E. triglochidiatus E. engelmannii 23 2 (90-270 degrees) 24 33 Fisher Exact Probability Test, p < 0.001. 356 Great Basin Naturalist Vol. 42, No. 3 Discussion Felger and Lowe (1967) described changes in the form of Lophocereiis schottii over a latitudinal gradient, with a tendency for larger diameter and a reduced number of stem ribs toward the colder, northern edges of its distribution. Also Niering, Whittaker, and Lowe (1963) demonstrated an increase in the diameter of Carnegiea gigantea at the northern limits of its distribution. These changes have the effect of reducing the sur- face-to-volume ratio and increasing the time lag before the tissues suffer damages from freezing temperatures, as has been simulated by Lewis and Nobel (1977) and Nobel (1978). These changes occur within species, but the results may be extrapolated to those observed for the two species of Echinocereus. In form, E. triglochidiatus has a more closed canopy than does E. engehnannii, because its stems grow in contact with one another. Hence its surface-to-volume ratio is reduced, and the species approximates in form a solid cylinder or "barrel." Echinocereus trighchidiatus is found only at the juniper-pinyon site where exposure to freezing temperatures at night can be a severe problem. The time lag, in which the internal stem temperatures are still increasing because of conductance between stems after the sun sets, may enable E. trig- lochidiatus to survive low night temper- atures. In contrast, E. engehnannii may be unable to survive low night temperatures due to its more open growth form. As a result, it is only established at the colder juniper-pin- yon site on unshaded, south-facing rocky slopes. Here it can warm up rapidly in the morning, minimizing the time during which its tissues are exposed to freezing temper- atures. Its growth form at the juniper-pinyon site, rather than approximating the com- pactness of E. triglochidiatus, is open due to the hot summer temperatures experienced in Table 3. Shade utilization by individuals of Echino- cereus triglochidiatus and E. engehnannii in the juniper- pinyon zone of the Providence Mountains, California. Species Shade No shade E. triglochidiatus E. engehnannii 67 5 5 38 x2 = 72.8 p < 0.01. this microhabitat. The smaller maximum size, attained by E. engehnannii at the juniper- pinyon site, is probably the result of its estab- lishment at the extreme upper limits of its elevational range, where its growth rate is slower and less constant and its probability of survival to the maximum sizes attained at the desert site is very low. At the desert site, freezing temperatures are less of a problem. High summer temperatures appear to be more critical there. One way of reducing the heat load of an individual is to open the can- opy, permitting convective cooling as air moves through the canopy. Although morphology appears to be an adaptation to the extremes in temperature that Echinocereus experiences, each species must cope with problems posed when the op- posite climatic conditions occur. For ex- ample, hot summer days do occur at the juni- per-pinyon site and freezing winter nights do occur at the desert site, although at a reduc- ed frequency in comparison with their oppo- site extremes. Microhabitat differences be- come important in moderating the heat load and exposure to freezing temperatures. Dur- ing the hot summer months, when the sun is directly overhead at midday, E. triglochi- diatus is usually in the shade. Its daily heat load is reduced, because ambient temper- atures in the shade are lower (Bannister 1976, Yeaton et al., in review). Conversely, in the winter months, when the sun is at a lower angle, E. triglochidiatus may be in direct sun- light during part of the day. At the desert site, E. engelmannii may avoid the effects of freezing temperatures by using unshaded mi- crohabitats in which individual stems can heat rapidly in the early morning as the sun- light strikes them. One comment should be made about the temperature measurements made. The great difference in surface temperature, recorded in the morning hours, is the direct result of differences in the canopy structure for the two species. As a result of its more open can- opy, the stems of E. engehnannii are exposed at times to direct sunlight and heat more rap- idly than do the stem surfaces of the closed- canopied E. trigiochidiatus, which are always shaded. For this reason surface and internal temperatures of E. triglochidiatus fluctuate similarly over the course of the day, and September 1982 Yeaton: Cactus Habitat Utilization 357 o. 30 0) t- J 25 o k- E < 0) a E 20 H- 15 0) ^-» 0 a E .0) c 0) !5 E < E o 0) o c 0) 0900 1200 1500 1800 Time 2100 2400 Fii;. 1. The relationship between ambient temperature and surface and internal stem temperatures for trans- planted Echinocereiis engelmannii and E. triglochidiatus in an open site over a 15-hour period on 15 September 1980. (a). Ambient air temperatures; (b), The differences from ambient temperature for surface ( ), 6 cm deep ( ), and 10 cm deep ( ) probes in or on the central stem of E. engelmannii; (c), the same as b. but for E. triglochidiatus. 358 Great Basin Naturalist Vol. 42, No. 3 Table 4. Means and standard errors for size measurements of the 10 largest individuals of Echinocereus triglochi- diatus and E. engehnannii found in each habitat utilized in the juniper-pinyon and desert zones of Providence Moun- tains, Cahfomia. Habitat Diameter (em) No. Stems No. Stems/Diameter Individual Stem Diameter (mm) Juniper-pinyon £. triglochidiattis- E. engelmwinii - Desert E. engehnannii - -Rocky -Pinyon -Wash -Rocky -Rocky 45.20 ± 3.60 39.50 ± 4.50 34.50 ± 2.41 19.20 ± 1.69 47.20 ± 4.30 86.40 ± 7.44 76.40 ± 11.55 47.60 ± 4.43 10.00 ± 0.67 .30.30 ± 3.11 1.93 ± 0.10 1.77 ± 0.17 1.41 ± 0.11 0.55 ± 0.05 0.67 ± 0.07 5.18 ± 0.16 4.82 ± 0.15 widely divergent temperatures are recorded in the same period for E. engelmannii. The difference in surface temperature is further exacerbated by the differential movement of air through the canopies of the two species, resulting in different rates of convective cool- ing. Thus, surface temperatures may not be compared with the internal stem temper- atures except as general trends. This is be- cause the temperatures recorded on the sur- face are a direct response to environmental conditions, but the internal stem temper- atures represent various degrees of inte- gration of these same environmental condi- tions. The internal stem temperatures may be compared between the 6 cm and 10 cm depths and, as would be expected, the 10 cm depth becomes warmer than the 6 cm depth as the day progresses. I have concentrated in this study on the differences in form of two species of Echino- cereus. Other differences such as spine cov- erage, spine color, apical pubescence, and tis- sue thermal properties have been demonstrated as being important in the regu- lation of heat load in Cactaceae (Nobel 1978). Differences between the two species do exist for some of these characteristics (Benson 1969). Echinocereus engehnannii is more heavily spined and its spines are lighter colored than those of E. triglochidiatus. Also, the central spines of E. engelmannii are flat- tened in contrast to those of E. triglochi- diatus. Tliese factors contribute to its adapta- tion to imshaded microhabitats by increasing its albedo (Gibbs and Patten 1970, Nobel 1978). Additionally, E. triglochidiatus has permanent apical pubescence on mature stems, but E. engelmannii loses its pub- escence after 2-3 years. Apical pubescence may have an insulative effect (Nobel 1978), which may be important in survival of stems of E. triglochidiatus in cold habitats. I have no information on the thermal properties of stem tissues. Thus, the differences in morphology ob- served in Echinocereus engelmannii and E. triglochidiatus appear to be adaptations for avoidance of extreme climatic conditions in their preferred habitats, and differences in microhabitats enable the individual to avoid the opposite extremes should they occur. In other words, the constraints imposed by mor- phology are ameliorated by differences in mi- crohabitat use. Acknowledgments I thank R. W. Eckert, W. Work, and R. W. Yeaton for their assistance in this study. Literature Cited Bannister, P. 1976. Introduction to physiological plant ecology. Halstead Press, New York. Benson, L. 1969. The native cacti of California. Stanford Univ. Press, Stanford. Felger, R. W., ANn C. H. Lowe. 1967. Clinal variation in the surface-volume relationships of the colum- nar cactus Lophocereus schottii in northwestern Mexico. Ecology 48:5.30-536. Gates, D. M. 1962. Energy exchange in the biosphere. Harper and Row, New York. Gibbs, J. G., and D. T. Patten. 1970. Plant temper- atures and heat flux in a Sonoran Desert ecosys- tem. Oecologia 5:165-184. Lewis, D. A., and P. S. Nobel. 1977. Thermal energy exchange model and water loss of a barrel cactus, Fewcacttis acnnthodes. Plant Physiology 60:609-616. September 1982 Yeaton: Cactus Habitat Utilization 359 Mozingo, H. N., and P. L. Comanor. 1975. Implications of the thermal response of Ferocactus acanthodes. Supplement of the Cactus and Succulent J. (U.S.):22-28. NiERiNG, W. A., R. H. Whittaker, and C. H. Lowe. 1963. The saguaro: a population in relation to en- vironment. Science 142:15-23. Nobel, P. S. 1978. Surface temperatures of cacti: in- fluences of environmental and morphological fac- tors. Ecology 59:986-996. 1980a. Morphology, surface temperatures, and northern limits of columnar cacti in the Sonoran Desert. Ecology 61:1-7. 1980b. Interception of photosynthetically active radiation by cacti of different morphology. Oeco- logia 45:160-166. OosTiNG, H. J. 1956. The study of plant communities. W. H. Freeman and Co., San Francisco. RoDRiQUEZ, S. B., F. B. Perez, and D. D. Montenegro. 1976. Eficiencia fotosintetica del nopal (Opuntia spp.) en relacion con la orientacion de sus cla- dodes. Agrociencia 24:67-77. SiEGEL, S. 1956. Non-parametric statistics. McGraw-Hill, New York. Steel, R. G. D., and J. H. Torrie. 1960. Principles and procedures of statistics. McGraw-Hill, New York. Trombulak, S. C, and M. L. Cody. 1980. Elevational distribution of Pinus edulis and P. monophylla (Pinaceae) in the New York Mountains, eastern Mojave Desert, Madrono 27:61-67. Yeaton, R. I., and M. L. Cody. 1979. The distribution of cacti along environmental gradients in the Sonoran and Mojave deserts. J. Ecology 67:529-541. Yeaton, R. I., R. Karban, and H. Wagner. 1980. Mor- phological growth patterns of Saguaro {Carnegiea gigantea: Cactaceae) on flats and slopes in Organ Pipe Cactus National Monument, Arizona. Southwestern Nat. 25:339-349. Yeaton, R. I., E. B. Layendecker, K. S. Sly, and R. W. EcKERT. Some aspects of microhabitat differences between two species of cactus in the mixed Chaparral-Inland Coastal Sage Association of southern California. In review. FIRST SPECIMEN OF THE SPOTTED BAT (EUDERMA MACULATUM) FROM COLORADO Robert B. Finley, Jr.,' and James Creasy^ Abstract.— A spotted bat (Euderma maculatum) was picked up at the headquarters of Browns Park National Wildhfe Refuge, Moffat County, Colorado, on 29 August 1981. The spotted bat {Euderma maculatum) has not been reported from Colorado, although its presence in the state was considered prob- able by Armstrong (1972). The specimen re- ported here was an accidental find, like many others previously reported (Barbour and Davis, 1969). On 29 August 1981, at about 1000 MDT on a clear day. Creasy found an adult female spotted bat resting on a concrete slab behind the shop in the old headquarters area of the Browns Park National Wildlife Refuge, 1630 m, Moffat County, Colorado. When touched, it spread its wings, raised its head and ears, and opened its mouth, but made no effort to fly. It was prepared as a study skin, skull, and alcoholic carcass; no sign of injury or illness was found during preparation. It was depos- ited in the Biological Surveys collection at Fort Collins (BS/FC 7557). Browns Park is a wide, mountain-ringed valley or "park" about 25 km long, through which the Green River meanders before en- tering Gates of Lodore, a narrow gorge at the north end of Dinosaur National Monument. The headquarters site is 9 km north and 2 km west of Gates of Lodore, on a strip of cotton- wood bottom between the left bank of the Green River and Harry Hoy Bottom, a cat- tail-bulrush marsh. The nearest cliffs are on the steep rocky mountainside 3 km southwest across the Green River. Cliffs a few hundred meters high form the walls of the Gates of Lodore 10 km to the south. The rocky moun- tainsides encircling Browns Park are wooded with pinyon-juniper. The nearest locality previously reported for the spotted bat was Salt Lake City, Utah, approximately 255 km to the west (Durrant 1935). Nearest known localities in other di- rections are 5 mi NW Monticello, Utah, ap- proximately 355 km to the south (Benson 1954); and Byron, Bighorn Co., Wyoming, approximately 440 km to the north (Mickey 1961). External measurements (in millimeters) of the specimen from Browns Park NWR are: total length, 119; length of tail, 53; length of hind foot, 13; length of ear, 42; length of tragus, 15; width of tragus, 5; length of fore- arm, 52; and weight, 13.5 g. Cranial measure- ments (in mm) taken as described by Handley (1959) are: greatest length of skull, 19.4; zygomatic breadth, 10.7; interorbital breadth, 4.2; breadth of braincase, 9.4; depth of braincase, 5.7; maxillary toothrow, 6.3; postpalatal length, 8.1; and palatal breadth, 7.2. Literature Cited Armstrong, D. M. 1972. Distribution of mammals in Colorado. Mono. Mus. Nat. Hist., Univ. of Kans. 3. 415 pp. Barbour, R. W., and W. H. Davis. 1969. Bats of Ameri- ca. Univ. Press of Kentucky. 286 pp. Benson, S. B. 1954. Records of the spotted bat (Euderma mocidata) from California and Utah. J. Mamm. .35(1): 117. Durrant, S. D. 1935. Occurrence of the spotted bat in Utah. J. Mamm. 16(3):226. Handley, C. O., Jr. 1959. A revision of the American bats of the genera Euderma and Plecotus. Proc. U.S. Natl. Mus. 110:95-246. Mickey, A. B. 1961. Record of the spotted bat from Wyoming. J. Mamm. 42(3):401. 'U.S. Fish and Wildlife Service, 1300 Blue Spruce Drive, Fort Collins, Colorado 80524-2098. 'Browns Park National Wildlife Refuge, Greystone Route, Maybell, Colorado 81640. 360 STATUS OF INTRODUCED FISHES IN CERTAIN SPRING SYSTEMS IN SOUTHERN NEVADA Walter R. Courtenay, Jr.," and James E. Deacon^ .Abstract.— We record eight species of exotic fishes as established, reproducing populations in certain springs in Clark, Lincoln, and Nye counties. Nevada. These include an unidentified species of Hi/postomus, Cyprinus carpio, Poecilia mexicana. Poecilia reticulata, a Xiphophorus hybrid, and Cichlasoma nigrofasciatiim. Tilapia mariae, estab- lished in a spring near the Overton Arm of Lake Mead, and Tilapia zilli, established in a golf course pond in Pah- rump Valley, are recorded for the first time from Nevada waters. Though populations of transplanted Gambusia af- finis persist, other populations of Poecilia latipinna are apparently no longer extant. Cichlasoma severum, Notemigomis crysoleucas, Poecilia latipinna, and Carassius auratus were apparently eradicated from Rogers Spring in 1963. Miller and Alcorn (1943), Miller (1961), La Rivers (1962), Deacon et al. (1964), Hubbs and Deacon (1964), Minckley and Deacon (1968), Minckley (1973), Hubbs et al. (1974), Deacon (1979), Hardy (1980), and others re- corded the presence of non-native fishes in Nevada. In those papers, it was stressed that the introduction of nonnative fishes, be they exotic (of foreign origin) or transplants native to otlier areas of the United States, can have serious, adverse impacts on the depauperate and often highly endemic fish fauna in the southwestern U.S. Deacon et al. (1964) em- phasized that most of the endemic fishes are small and, therefore, subject to more adverse impacts through introductions of small bait or ornamental fishes than with earlier in- troductions of larger fishes, a subject re- viewed by Hubbs and Broderick (1963). In this paper, we document eight species of exotic fishes in three counties of southern Nevada. One of these fishes, the spotted ti- lapia {Tilapia mariae), was previously known to have become established only in Florida (Hogg 1974, 1976, Courtenay and Robins 1975, Courtenay 1979b, 1980, Courtenay and Hensley 1979, 1980). Another, the redbelly tilapia {Tilapia zilli), has been recorded as es- tablished in Arizona, California, and Texas (Courtenay and Hensley 1980). Our purpose is to update the status of introduced fishes in southern Nevada, primarily from the reports of Deacon et al. (1964) and Hubbs and Dea- con (1964). Clark County Indian Spring is 2 km south of U.S. High- way 95, approximately 62 km northwest of Las Vegas in the village of Indian Springs. Minckley (1973) recorded a suckermouth cat- fish {Hypostomiis) as successfully established since at least 1966 "in a warm spring in southern Nevada"; this reference was to In- dian Spring. Brief examination of Indian Spring by Deacon .since 1966 demonstrates that the population has remained common and continues to reproduce successfully. The deeply imdercut banks with numerous tree roots and holes provide excellent refuge for this rather cryptic species. Our collection on 18 October 1980 yielded only one specimen, despite repeated seining. A local resident ad- vised us that local youngsters had removed as many as 500 individuals in recent months, probably for sale to pet shops in Las Vegas; this, predation on eggs by an introduced snail {Melanoides tiiberciilata), competition from other introduced fishes, or a combination of these factors could explain the apparent pop- ulation decline of Hypostomus he felt had oc- curred. On 25 June 1981 a reexamination of the spring pond using a face mask and a light to observe the undercut banks demonstrated 'Department of Biological Sciences, Florida Atlantic University, Boca Raton. Florida 33431. 'Department of Biology, University of Nevada, Las Vegas, Nevada 891.54 361 362 Great Basin Naturalist Vol. 42, No. 3 that Hypostomus continues to exist in rela- tively high density in Indian Spring. Individ- uals of all sizes were seen hiding in or around almost every available hole, tree root, or rock. Suckermouth catfish have been recorded as established in Texas (Barron 1964, Hubbs et al. 1978) and Florida (Courtenay and Robins 1973, Courtenay 1979b, 1980). The specimen collected at Indian Spring is morphologically distinct from those collected in Texas and Florida and represents a third species of Hy- postomus established in the U.S. Deacon (pers. comm. to D. A. Hensley) also recorded the green swordtail (Xiph- ophonis helleri) from Indian Spring in 1975. Although we found no green swordtails at In- dian Spring on 18 October 1980, a yellow to pale orange hybrid swordtail (probably X. helleri x X. maculatus) was found to be the dominant fish in number of individuals. Gup- pies (Poecilia reticulata) were also abundant and several large common carp (Cyprinus carpio) were seen but not collected. The only other species present was the mosquitofish {Gambusia af finis). Blue Point Spring is in the Lake Mead Na- tional Recreational Area above the Overton Arm of Lake Mead, approximately 68 km northeast of Las Vegas. Deacon et al. (1964) recorded guppies, shortfin mollies {Poecilia mexicana), and the southern platyfish {Xiph- ophorus maculatus) from Blue Point Spring. In a collection made 18 October 1980, no guppies or southern platyfish were found; however, shortfin mollies, convict cichlids, and a single spotted tilapia with damaged pelvic fins were captured. Deacon et al. (1964) also reported a transplant, the sailfin molly {Poecilia latipinna), from Blue Point Spring; we did not collect this species there and it is assumed that the population died out. Rogers Spring is located 2 km southwest of Blue Point Spring in the Lake Mead National Recreational Area. Deacon et al. (1964) re- ported convict cichlids, goldfish {Carassius auratus) and transplanted sailfin mollies, golden shiners {Notemigonus crysoleucas), and mosquitofish from Rogers Spring. Hubbs and Deacon (1964) reported convict cichlids, shortfin mollies, transplanted sailfin mollies, and the banded cichlid {Cichlasoma severum) from this spring. An attempt to remove the introduced fishes was made in December 1963. We seined Rogers Spring on 18 Octo- ber 1980. The dominant fish (in numbers and biomass) was the spotted tilapia {Tilapia mariae). Shortfin mollies were very abundant, convict cichlids were rare, and two guppies were collected. Underwater observations made prior to the seine hauls correlated well with the population densities revealed by seining. No banded cichlids, sailfin mollies, golden shiners, or mosquitofish were seen or collected, and they are considered as no long- er extant in Rogers Spring. Of particular interest were the underwater observations on spotted tilapia prior to sein- ing activities. Most of the tilapias, even large individuals (above 100 mm SL), displayed the banded juvenile pattern as illustrated by Thys van den Audenaerde (1966). Only a relatively few large individuals showed the typical adult pattern of spots on their sides (Fig. 1). Moreover, it was later noted, following sein- ing, that most of the tilapias displaying the juvenile color pattern were missing parts or most of their pelvic fins (Fig. 2) and that other fins (particularly the soft dorsal and up- per half of the caudal) often showed damage. The only fish in Rogers Spring capable of in- flicting such damage were the adults with the typical adult color pattern; none of those had damaged fins. At no time have population densities of Ti- lapia mariae in Florida been observed to be approaching those seen in Rogers Spring. It is probable that the trophic and spatial car- rying capacities of Rogers Spring for spotted tilapia have been reached and that this be- havioral hierarchy, led by a few highly ag- gressive individuals, has developed to control further overpopulation. Lincoln Gounty Deacon et al. (1964) did not find any exotic fishes at either Grystal Spring or Ash Springs in the Pahranagat Valley, some 145 km north-northwest of Rogers Spring during col- lecting trips on 2 February and 9 March 1963. They did find an introduced population of mosquitofish at Ash Springs that was re- ported earlier by Miller and Hubbs (1960). During a collecting trip on 3 June 1964, September 1982 COURTENAY, DeACON: INTRODUCED NevADA FiSHES 363 FiiT. 1. An advilt spotted tilapia. 205 mm standard length. Drawing by Francis McKittrick Watkins. convict cichlids, shortfin mollies, and trans- planted sailfin mollies were discovered in Ash Springs (Hubbs and Deacon 1964). The prob- able source of these introduced fishes was suggested as Rogers Spring because all three species were known to exist there. The same three non-native fishes subsequently spread from Ash Springs to Crystal Spring, 8 km to the north-northwest. On 17 October 1980, surface and under- water observations, as well as seine collec- tions, made in the outflow of Ash Springs showed convict cichlids to be dominant in biomass, followed by shortfin mollies that were dominant in numbers of individuals. Transplanted mosquitofish were common and no sailfin mollies were seen or collected. In- troduced fishes were somewhat uncommon in a pool occupied by 35-40 individuals of the Pahranagat roundtail chub {Gila robusta prdani). Nye County Goldfish, Carassius auratus, were reported by Deacon et al. (1964) and by Deacon (1979) in Pahrump Valley at Manse Ranch Spring. Manse Ranch Spring dried up for a short time in the summer of 1975 (Soltz and Naiman 1978), thus eliminating the goldfish. About 10 km to the north of Manse Ranch Spring is the site of Pahrump Spring, origi- nally the largest spring in Pahrump Valley. Pahrump Spring failed in the late 1950s be- cause of excess pumping of groundwater for irrigation. During the late 1970s the sur- rounding area was subjected to a land devel- opment plan that included construction of a golf course and park area. The ponds and small stream associated with this devel- opment use groundwater from the vicinity of the former spring source. Examination of the main pond in Cottonwood Park by the Ne- vada Department of Wildlife on 4 March 1980 and by one of us (Deacon) on 16 July 1981 revealed the presence of large numbers of goldfish and mosquitofish and relatively small numbers of the redbelly tilapia, Tilapia zilli. The specimens of redbelly tilapia col- lected were all small, but one pair of larger fish (to ca 250 mm) was seen in the water in July, apparently guarding nests or young. The manager reported that large numbers of Tilapia died last winter when the water tem- perature reached 58 F (14 C). This species has maintained itself through at least two 364 Great Basin Naturalist Vol. 42, No. 3 Fig. 2. A spotted tilapia, 73.5 mm SL, from Blue Point Spring. Note damaged pelvic fins. winters and has reproduced during the sum- mers of 1980 and 1981. A small population is able to exist through the winter apparently because the inflowing water maintains a tem- perature of about 70 F (21 C). Discussion Courtenay et al. (1974) termed the promis- cuous introduction of exotic fishes to new en- vironments as biological pollution. Unlike chemical or thermal pollutants, biological ad- ditives have the ability to reproduce and ex- pand their ranges. Moreover, many exotic fishes may appear to be trophic specialists in their native range but prove to be passive generahsts (Birkeland and Neudecker 1981) in new environments where most or all of the biological constraints of the native range are absent (Courtenay and Hensley 1980). Such trophic adaptation, coupled frequently with equally adaptable behavioral traits, permits the exotic species to disrupt habitats and niches {sensu lato) in new environments. The introduction of any non-native fish will result in alterations in the host ecosys- tem. Such alterations may range from minor, almost unnoticeable changes in native fish populations to the extinction of one or more native species (Courtenay 1979a, Courtenay and Hensley 1980). In Florida, for example, comparatively small ornamental fishes, cich- lids in particular, are able to successfully in- vade and become dominant in waters that contain larger native piscivores such as large- mouth bass {Micropterus salmoides) and Flor- ida gar (Lepisosteus platyrhinciis). Invasion and domination by nonnative fishes in many parts of the desert southwest is facilitated by the absence of piscivores and the presence of small endemic fishes that have had no pre- vious experience with trophic passive gener- ahsts or piscivores. Therefore, a fish that feeds on phytoplankton or detritus in its September 1982 COURTENAY, DeACON: INTRODUCED NeVADA FiSHES 365 native range and either becomes a predator of smaller fishes or feeds on algae in the new environment is a clear threat to many native fishes in the southwestern U.S. It is obvious that there have been several changes in the introduced fish fauna in south- em Nevada since 1963. Species compositions have changed. The green swordtail popu- lation in Indian Spring is gone and has been replaced with a hybrid of Xiphophorus; a population of a suckermouth catfish remains extant there. Guppies, southern platyfish, and sailfin mollies appear to be absent in Blue Point Spring, and at least one spotted tilapia was released, probably recently, before our collections there. Sailfin mollies, mosquito- fish, and the banded cichlid are absent in Rogers Spring, the latter probably due to the eradication effort in 1963; convict cichlids are now rare, but guppies and spotted tilapia (the latter now dominant) have been added. Sailfin mollies appear to be absent from the outflow of Ash Springs in the Pahranagat Valley, whereas convict cichlids and shortfin mollies are dominant, 17 years after their introduction. Frequent and closer monitoring of in- troduced fish populations, particularly in areas that are potential or recognized release sites for unwanted pet fishes in southern Ne- vada and other areas in the desert southwest, is needed. These "pockets" of introduced fishes serve as potential and probable sources for futiu-e introductions elsewhere as appar- ently occurred with the transfer of convict cichlids and shortfin mollies from Rogers Spring to Ash Springs in 1963. A further in- dication of the constant or continuing nature of this problem is a verbal report from Mr. Charles Orr, a Las Vegas member of the American Cichlid Association, that he saw and identified several specimens of the "mar- malade" form of Pseudotropheus zebra when he visited Rogers Spring in July 1981. Although the disappearance of some in- troduced fishes in southern Nevada in recent years may be of some interest, it is far more important tliat studies be initiated to define the impacts of introduced species on native fishes. Species interactions between non- native and native fishes have been suggested as reasons for declines in populations of na- tive fishes (Deacon et al. 1964, Hubbs and Deacon 1964, Courtenay and Hensley 1980), but to date the exact mechanisms for these declines have not been examined. The "cause and effect" for such declines is suggested strongly but requires in situ observations and evaluations. Finding an established population of spotted tilapia in southern Nevada is particu- larly disturbing. Although this fish is omniv- orous, it shows a preference for green algae in Florida and in its native range has been described as "une forme intermediaire entre les Tilapia herbivores et les especes micro- phages" (Thys van den Audenaerde 1966). The bottom of Rogers Spring was devoid of green algae, doubtlessly due to grazing by spotted tilapia. A trophic preference of this type, coupled with omnivory, could prove disastrous to several endemic species and sub- species of southwestern fishes if this western African cichlid were moved elsewhere. We therefore recommend its immediate eradica- tion. Much the same result could be expected by future introductions of other tilapias {sen- sii lato) or such seemingly harmless fishes as Hypostomus spp. Also, the potential in- troduction of fish parasites via exotic fish vectors exists (Lachner et al. 1970, Courtenay and Robins 1975, Courtenay 1979a) and has been suggested to have occurred in southern Nevada (Deacon 1979) and elsewhere (Hoff- man 1970, Bauer and Hoffman 1976). Fish introductions pose as great a tlireat to the continued existence of the depauperate fish fauna in southern Nevada and adjoining states as does water withdrawal for agricul- tural, domestic, military, and industrial uses or other habitat modifications. Acknowledgments We thank Paul Greger and Thom Hardy for assistance in the field and James A. McCann, Robert R. Miller, and Jeffrey N. Taylor for their helpful comments on the manuscript. This study was supported by Contract 14-16-0009-78-021, Cooperative Agreement 14-16-0009-80-952, and an Inter- governmental Personnel Act Assignment from the Department of the Interior, U.S. Fish and Wildlife Service, and by sabbatical fmiding from Florida Atlantic University. 366 Great Basin Naturalist Vol. 42, No. 3 Literature Cited Barron, J. C. 1964. Reproduction and apparent over- winter survival of suckermouth armored catfish, Plecostomus sp., in the headwaters of the San An- tonio River. Texas J. Sci. 16:449-450. Bauer, O. N., and G. L. Hoffman. 1976. Helminth range extension by translocation of fish. Pages 163-172 in L. A. Page, ed., Wildlife diseases. Ple- num Publ. Corp., New York. BiRKELAND, C, AND S. Neudecker. 1981. Foraging be- havior in two Caribbean chaetodontids: Chaeto- don capistratiis and C. aculeatus. Copeia 1981 (1):169-178. CouRTENAY, W. R., Jr. 1979a. The introduction of exotic organisms. Pages 237-252 in H. P. Brokaw, ed.. Wildlife and America. Government Printing Of- fice, Washington. 1979b. Biological impacts of introduced species and management policy in Florida. Pages 237-257 in R. Mann, ed., Exotic species in mari- culture. MIT Press, Cambridge, Massachusetts. 1980. Exotic fish: environmental roulette. Water %)ectrum 12(4): 10-17. CouRTENAY, W. R., Jr., AND D. A. Hensley. 1979. Range expansion in southern Florida of the introduced spotted tilapia. Environ. Conserv. 6(2): 149-151. 1980. Special problems associated with mon- itoring exotic species. Pages 281-307 in C. H. Hocutt and J. R. Stauffer, Jr., eds.. Biological monitoring of fish. Lexington Books, Lexington, Massachusetts. CouRTENAY, W. R., Jr., and C. R. Robins. 1973. Exotic aquatic organisms in Florida with emphasis on fishes: a review and recommendations. Trans. Amer. Fish. Soc. 102(1):1-12. 1975. Exotic organisms: an unsolved, complex problem. BioScience 25(5):306-313. CouRTENAY, W. R., Jr., H. F. Sahlman, W. W. Miley II, and D. J. Herrema. 1974. Exotic fishes in fresh and brackish waters of Florida. Biol. Con- serv. 6(4):292-,302. Deacon, J. E. 1979. Endangered and threatened fishes of the west. Great Basin Nat. Mem. No. 3:41-64. Deacon, J. E., C. Hubbs, and B. J. Zahuranec. 1964. Some effects of introduced fishes on the native fish fauna of southern Nevada. Copeia 1964 (2):384-388. Hardy, T. 1980. The Inter-Basin area report. Proc. Desert Fish. Coun. 11:5-21. Hoffman, G. L. 1970. Intercontinental and trans- continental dissemination and transfaunation of fish parasites with emphasis on whirling disease {Myxosoma cerebralis). Pages 69-81 in S. F. Snieszko, ed., A symposium on diseases of fishes and shellfishes. Amer. Fish. Soc. Spec. Publ. 5. Hogg, R. G. 1974. Environmental hazards posed by exotic fish species newly established in Florida. Environ. Conserv. 1(3): 176. 1976. Established exotic cichlid fishes in Dade County, Florida. Florida Sci. 39(2):97-103. Hubbs, C, and H. J. Broderick. 1963. Current abun- dance of Gambusia gaigei, an endangered fish species. Southwest Nat. 8:46-48. Hubbs, C, and J. E. Deacon. 1965. Additional in- troductions of tropical fishes into southern Ne- vada. Southwest Nat. 9(4):249-251. Hubbs, C, T. Lucier, G. P. Garrett, R. J. Edwards, S. M. Dean, E. Marsh, and D. Belk. 1978. Survi- val and abimdance of introduced fishes near San Antonio. Texas J. Sci. 30(4):369-376. Hubbs, C. L., R. R. Miller, and L. C. Hubbs. 1974. Hy- drographic history and relict fishes of the north central Great Basin. California Acad. Sci. Mem. 7:1-259. Lachner, E. a., C. R. Robins, and W. R. Courtenay, Jr. 1970. Exotic fishes and other aquatic organ- isms introduced into North America. Smithson. Stud. Zool. 59:1-29. La Rivers, I. 1962. Fishes and fisheries of Nevada. Ne- vada St. Fish and Game Comm. 782 pp. Miller, R. R. 1961. Man and the changing fish fauna of the American southwest. Pap. Mich. Acad. Sci., Arts, and Lett. 46:365-404. Miller, R. R., and J. R. Alcorn. 1946. The introduced fishes of Nevada, with a history of their in- troduction. Trans. Amer. Fish. Soc. 73:U.3-193. Miller, R. R., and C. L. Hubbs. 1960. The spiny-rayed cyprinid fishes (Plagopterini) of the Colorado River system. Misc. Publ. Mus. Zool. Univ. Mich- igan 115:1-39. MiNCKLEY, W. L. 1973. Fishes of Arizona. Arizona Game and Fish Dept., Phoenix. 293 pp. MiNCKLEY, W. L., AND J. E. Deacon. 1968. South- western fishes and the enigma of "endangered species." Science 159:1424-1432. SoLTZ, D. L., AND R. J. Naiman. 1978. The natural his- tory of native fishes in the Death Valley system. Natural History Museum of Los Angeles County, Science Series 30:1-76. Thys VAN den Audenaerde, D. F. E. 1966. Les tilapia (Pisces, Cichlidae) du sud-Cameroun et du Ga- bon; etude systematique. Ann. Mus. Roy. Afr. Cent., ser. 8, Zool. 153:1-98. Wilson, B. L., J. E. Deacon, and W. G. Bradley. 1966. Parasitism in the fishes of the Moapa River, Clark County, Nevada. Trans. California-Nevada Sec. Wildl. Soc. pp. 12-23. A NEW SPECIES OF PENSTEMON (SCROPHULARIACEAE) FROM THE UINTA BASIN OF UTAH AND COLORADO John Larry England' Abstract.- Named as a new species is Penstemon albifluvis ]. L. England. The species is known from the Uinta Basin of Utali and Colorado. The Uinta Basin of Utah and Colorado har- bors numerous endemic species of plants, with many of these restricted to the Green River Formation. It is not surprising that this habitat should give rise to yet another nar- rowly restricted species. The author, while employed by the Bureau of Land Manage- ment, encountered a novel Penste7non grow- ing on oil shale ledges of the Green River Formation immediately adjacent to the White River. Subsequent examination re- vealed this botanical novelty to be distinct from any described taxon. Hence, it is de- scribed herein. Penstemon albifluvis England, sp. nov. Ab Penstemon scariosus et P. strictus in calycibus et corollis brevioribus et foliis radi- calibus paucis vel nullis et pubis antheris bre- vioribus differt. Perennial herb; stems ascending to erect, (1) 1.5-4.5 (5) dm tall, (1) 5-20 clustered on a frequently branched caudex, surmounting a taproot; herbage glabrous; leaves entire or with crisped margins (2) 4-10 (12) cm long, 4-6 (11) mm wide, lacking basal leaves or these poorly developed, lower cauline leaves narrowly oblanceolate, petiolate, the upper cauline ones mostly linear to narrowly lan- ceolate, sessile, often crisped margined; thyrse secvmd, of 3-10 verticillasters, the up- per leafy bracts much reduced, the cymes (1) 2- to> 4-flowered, the axis, peduncles, and pedicels glandular-pubescent; sepals 4-6 (7) mm long, lanceolate, acute, glandular- pubescent, the margins narrowly scarious; co- rolla (18) 20-22 (24) mm long, gradually and broadly ventricose ampliate, the throat 6-7 mm broad, the tube 14-15 mm long, bila- biate, the upper lip projecting, arched, 7-8 mm long, the lobes of the lower lip spread- ing, pale lavendar, the lobes sometimes light blue, sparsely glandular-pubescent externally, the palate with two ridges 1 mm high on ei- ther side of the staminode, glabrous; stami- node 9-10 mm long, ending ca 1 mm short of the groove in the palate, straight apically, sparsely orange bearded, with hairs 0.2-.3 mm long, gradually enlarged apically, from 0.3-0.7 mm broad; fertile stamens reaching the orifice, the anther sacs 3-5 mm long, 0.5 mm broad, pubescent, the hairs white, 0.5 mm long or less, dehiscing the full length, black; capsule 8-11 mm long, broadly ovoid, acuminate; seeds ca 2 mm long. Type.- USA. Utah: Uintah Co., North bank of White River, ca one mile upstream from the Ignatio bridge, 2.5 miles airline dis- tance south of Bonanza, TIOS, R24E, SI, ele- vation 4975 ft, on raw shale slopes of the Evacuation Creek Member of the Green Riv- er Formation, associated with Eriogonum ephedroides, Cirsium barnebyi, Machaeran- thera grindelioides, Oryzopsis hymenoides, and Forsellesia meionandra, 9 June 1980, J. L. England 2046 (Holotype BRY; Isotypes to be distributed). The White River penstemon grows on raw shales with little soil development with Eriogonum, Cirsium, and Forsellesia in the desert saltbrush-sagebrush zone. It is endemic to east central Uintah County, Utah, and ad- jacent Rio Blanco County, Colorado. The plants flower from late May through June. 'U.S. Fish and Wildlife Service. 125 South State Street, Salt Lake City, Utah 84138. 367 368 Great Basin Naturalist Vol. 42, No. 3 Table 1. Characteristics oi Penstemon albifliivis and its relatives. Calyx length Corolla length Basal leaves Anther pubescence Habitat P. albifluvis P. scarious 4-6(7) mm (18) 20-22 (24) mm lacking or poorly developed hairs less than width of anthers (4) 6-9 mm (20) 24-30 mm well developed many hairs longer than width of anthers On poorly developed soils of higher elevations to deeper Green River Formation on well-developed soils at very xeric sites; lower elevations on variable 1500-1700 m geologic substrates 2000-3000 m P. strictus 3-8 (10) mm (20) 24-30 mm well developed many hairs longer than length of anthers higher elevations to deeper well-developed soils at lower elevations on variable geologic substrates 1800-3200 m Penstemon albifluvis is a member of the section Glabri (Pennell 1920) and has strong affinities to the pubescent anthered members of that section, most notably Penstemon sca- riosus and P. strictus. The characteristics of the White River penstemon and its relatives are compared in Table 1. Literature Cited Pennell, F. W. 1920. Scrophulariaceae of the Central Rockv Mountain States. Contr. U.S. National Herb! 20: 313-381. PHALACROPSIS DISPAR (COLEOPTERA: PHALACRIDAE), AN ELEMENT IN THE NATURAL CONTROL OF NATIVE PINE STEM RUST FUNGI IN THE WESTERN UNITED STATES' David L. Nelson- Abstract.— Larvae of the phalacrid beetle Phalacropsis dispar (LeConte) consumed aeciospores and the under- lying sporogenous mycehum, thereby destroying the aecia of all native western pine stem rust fiuigi studied. Aecia of the introduced white pine blister rust fimgus (Cronartium rihicola) were not found to be infested by the beetle. A close, if not obligate, biosis of the beetle apparently exists with the native rust fimgi, and their geographic distribu- tions closely coincide. Laboratory tests and field observations indicate that the beetle completes its life cycle in 30 to 40 days and apparently overwinters as an adult. Quantitative data on aeciospore inoculum destruction were beyond the means of this study; however, observations over a 12-year period evidenced widespread and extensive destruction of aeciospores. The beetle may be an effective element in the natural control of native pine stem rust fimgi. Natural control by secondary organisms could significantly reduce the selective pressure for high host resistance in a natural- ly evolving host-parasite population. Direct control of pine stem rust disease problems through application of chemicals and eradication of alternate hosts is usually expensive and of questionable effectiveness (Toko et al. 1967, Leaphart and Wicker 1968, Peterson and Jewel 1968, Carlson 1978). Various silvicultural methods (Peter- son 1966, Van Arsdel 1961, Nighswander and Patton 1965, Krebill 1968) and selection for host resistance appear to be the most accept- able means to immediate and long-term con- trol (Hanover 1966, Bingham et al. 1971, Hi- ratsuka and Powell 1976). Research on pine stem rust in the western United States has concentrated on the exotic white pine blister rust {Cronartium rihicola Fisch. in Rabenh.). The native rusts are currently not considered important enough to justify more than limit- ed study. Natural control other than host re- sistance may be an important factor in allow- ing this option. Answers for low-cost control of conifer rust diseases considered important could come from study of the natural control of native rust fungi. This may be especially true in the less extensively managed forests of western North America. Host tissue affected by pine stem rusts and the fruiting bodies of the aecial state of the rust fungi provide an attractive habitat and food source for many other organisms. These are mainly fungi (Mielke 1933, WoUenweber 1934, Powell 1971a,b,c, Byler et al. 1972b, Williams 1972, Kuhlman and Miller 1976, Kuhlman et al. 1976, Hiratsuka et al. 1979, Tsuneda and Hiratsuka 1979, 1980, Tsuneda et al. 1980, Kuhlman 1981a,b), insects (Snell 1919, Myren 1964, Coulson and Franklin 1970, Powell 1971d,e, Furniss et al. 1972, Powell et al. 1972, Kuhlman 1981b), and, to a lesser extent, rodents (Mielke 1935, Powell 1974) and mollusks (Hunt 1978). They may play a more important role than is recog- nized in the selection and limitation of these rust fungi through reduction of aecial inoculum. This study provides information on the identity, life history, and distribution of an aeciospore-consuming phalacrid beetle and suggests the possible importance of its de- struction of aecia and the activity of other re- ported secondary organisms in the natural control of some western native pine stem rust fungi. Review Many insects and other arthropods are as- sociated with pine stem and cone rusts in North America. Of special interest here are those that attack the rust fungi directly, are dependent upon them, consume large amounts of aeciospores, and consume or dam- age sporogenous mycelium. 'This article was written and prepared by a United States government employee on official time, and it is therefore in the public domain. 'Intermountain Forest and Range Experiment Station, Shrub Sciences Laboratory, Provo, Utah 84601. The initial observations for this study were made vhile the author was a student at the University of California, Berkeley. 369 370 Great Basin Naturalist Vol. 42, No. 3 Hubert (1923) described the activity of Epuraea ovata Horn (Coleoptera: Nitidu- lidae) larvae as consuming the entire aecio- spore mass and underlying stromatic myce- lium of the western gall rust fungus, Endocronartium harknessii (Peridermium harknessii). Powell (1971d) reviewed insects associated with pine stem rusts and listed 160 species of arthropods he found on Cronartium comandrae blister rust cankers on Pinus con- torta Dougl. Powell et al. (1972) listed those found associated with E. harknessii, C. com- andrae, C. coleosporioides, and C. comptoniae Arth. Using Graves's and Benick's system of classification, Powell (1971d) classed Epuraea obliquus Hatch, Paracacoxenus gluttatus Ha- ryd & Wheeler (Diptera: Drosophilidae), and a mite, Diapterobates principalis Berlese (Acarina: Ceratozetidae), as apparent true mycetobionts completely dependent on the aecia of C. comandrae for food. Epuraea ob- liquus was also found associated with C. co- leosporioides, Endocronartium harknessii, and C. comptoniae (Powell et al. 1972). Species of 12 genera of the insect orders Homoptera, Coleoptera, Lepidoptera, and Diptera associ- ated with comandra blister rust (C. com- andrae) cankers were considered to be appar- ent facultative mycetophiles. Several species of the mycetophilous type, especially the cone moth Dioryctria spp. (Lepidoptera: Pyralidae), extensively damaged areas of the aecial and spermogonial zone by burrowing in infected phloem tissue. Similar behavior of this moth has been observed on C. fusiforme Hedge. & Hunt (Coulson and Franklin 1970), C. strobilinum (Arth.) Peterson (Merkel 1958), C. comptoniae (Anderson and French 1964), £. harknessii (Byler et al. 1972a), and C. coleosporioides (Powell 197 Id). Powell (197 Id) regarded most of the 38 species of Hymenoptera he observed as parasitic on Lepidoptera, Diptera, and Coleoptera that inhabited the rust cankers. Furniss et al. (1972), examining cankers of white pine blis- ter rust in northern Idaho, found some of the species described by Powell (1971d) and Powell et al. (1972), but the number and di- versity of species were considerably less. The only apparent true mycetobiont type found by Furniss et al. was Paracacoxenus gluttatus. The extent to which pine stem rusts are damaged by these insects has received little study, and the impact of insect attacks on re- duction of aeciospore inoculum and on sub- sequent infection of pine has received even less. Over a seven-year period, between 32 and 57 percent of the C. comandrae cankers received obvious annual damage on 23 loca- tions, according to Powell (1971b). From 60 to 80 percent of the cankers were damaged yearly over this period. Insects infested up to 49 percent of recently dead or damaged galls of E. harknessii on Pinus radiata D. Don on one plot in California (Byler et al. 1972a). Nearly 45 percent of the E. harknessii galls studied by Wong (1972) in Manitoba and Sas- katchewan over a four-year period were mined by larvae of Dioryctria banksiella Mu- tuura, Munroe, and Ross, but Wong gave no information on damage to the rust fungus. Attacks by some insects that consume aecio- spores can be severe. Studies in southwestern Alberta (Powell 1971d) indicated a yearly 10 percent reduction in aeciospore production, with some years having a much higher per- centage. Epuraea obliquus occurred on as many as 80 percent of the sporulating can- kers on a single location, with at least 50 per- cent being attacked over a five-year period. Dipterous and lepidopterous larvae were ob- served on over 25 percent of sporulating can- kers in certain locations. The activity of Dio- ryctria sp., E. lengi Parsons, unknown dipterous larvae, and other insects apparently was of little importance in reducing aecio- spore inoculum of southern fusiform rust in North Carolina according to Kuhlman (1981b). Methods During other research on native western conifer rusts, a beetle was frequently ob- served in the aecia of pine stem rust fungi. The beetle's distribution and activity were studied on annual field trips from 1963 through 1975. Observations were made in coastal California from Fort Bragg south to the San Francisco Bay area and the Sierra Nevada, western Nevada from Lake Tahoe south to the Spring Mountains north of Las Vegas, northern Arizona south to Prescott, most of Utah, extreme western Wyoming, September 1982 Nelson: Natural Control of Pine Stem Rust 371 Fig. 1. Adult, pupal, and larval stages of Phalacropsis dispar, enlarged 14 times. southern Idaho from the Salmon River south, and east central Oregon. Because of the uncertain taxonomic status of some of the rust fungi involved and be- cause the study is concerned with the aecial state, the imperfect or peridermium name of some of the rust fimgi will be used for clarity throughout the remainder of the paper. To obtain information on beetle life his- tory, rust-infected stem sections that were also beetle infested were collected for labora- tory study. Infested rust samples were placed in one-liter glass beakers for observation. Drying of specimens was slowed by covering beaker tops with a plastic film punctured in several places to allow air exchange. Cheese- cloth was placed over the film to prevent es- cape of insects. Thus assembled, the speci- mens were held in the laboratory at normal room temperature, approximately 21 C. Both natural and fluorescent light was received during the day. Observations on development were made daily when possible. Dates when larvae matured, pupation began, and adult beetles emerged were recorded. Tliree locations were selected for annual field observation of beetle activity and rust development: (1) Bucks Lake, Plumas Na- tional Forest, Plumas Coimty, California; (2) Lee Vining Creek, Toiyabe National Forest, Inyo County, California; and (3) Red Canyon, Dixie National Forest, Garfield County, Utah. At location one, western gall rust, caused by Peridermium harknessii {Endocro nartiiim harknessii), was present on Pinus ponderosa Laws.; at location two, limb rust and stalactiform canker rust (both caused by Peridermium stalactiforme [Peterson 1968], an imperfect state of Cronartium coleospo- rioides), were present on Pinus jeffreyi Grev. & Balf. and P. contorta, respectively; and at location three, Powell limb rust (Peterson 1968), caused by Peridermium filamentosum, was present on Pinus ponderosa. Observa- tions were made at location one from 1963 through 1975 and at locations two and three from 1967 through 1975. These locations were visited one to three or more times each year during the aecial sporulation period. Results Habitat and Identification Numerous small, white-to-grayish beetle larvae (Fig. 1) were frequently found in the 372 Great Basin Naturalist Vol. 42, No. 3 Fig. 2. Beetle-damaged pine stem rusts (note weblike fecal debris over surface of stems): western gall rust (A) uno- pened, uninfested aecia (arrow) one-half actual size; (B) insect-damaged aecia; limb rust (C) unopened, uninfested aecia (arrow) actual size; (D) insect-damaged aecia (arrow points to unconsumed peridial fragments). aecial spore masses (Fig. 2A-D) of the vari- ous pine stem rust fungi observed. Their vo- racious consumption of aeciospores and of the underlying sporogenous mycelium effec- tively reduced aecia (Fig. 2A, C) to a mass of fecal debris within several weeks' time (Fig. 2B, D). Microscopic examination of young larvae revealed that the gut and in- September 1982 Nelson: Natural Control of Pine Stem Rust 373 testinal tract were full of aeciospores. Masses of spores were discharged in strings of fecal pellets. Several germination tests of aecios- pores from fecal pellets failed to show any vi- ability. Unconsumed spores in the same col- lections germinated. Spores of secondary fungi (such as Penicillitwi, which commonly invade aecial masses [Byler et al. 1972a]) were also consumed by the larvae. The per- idium of aecia was not consumed. Beetle specimens reared in the laboratory from larval-infested western gall and limb rust samples were used for identification. Larvae pupated and developed into shiny chestnut brown beetles about 3 mm in length (Fig. 1). They were identified as Phalacropsis dispar (LeConte) by Dr. Carl T. Parsons, who confirmed his identification by comparing specimens with the type specimen at the Mu- seum of Comparative Zoology, Harvard Uni- versity, Cambridge, Massachusetts. Phalacropsis is a monotypic genus de- scribed by T. L. Casey (1889-1890). The type specimen was collected in 1878 by E. A. Schwarz on a geological survey at 9400 ft on Veta Pass, Colorado, and described by T. L. Le Conte (1879). No host plant was men- tioned. Phalacropsis dispar apparently is rarely collected. A check of 20 insect mu- seums throughout the United States revealed only six collections as follows: California Academy of Sciences, Golden Gate Park- two collections, both from the Sierra Nevada, one on Piniis sp., no host listed for the other specimen; Ohio State University— two collec- tions, one from Yosemite National Park, Cali- fornia, the other from the Chiricahua Moun- tains of Arizona, no hosts were listed; and National Museum of Natural History— two collections, one from the Sitgreaves National Forest, Arizona, on Pinns ponderosa and the other from Pringle, South Dakota, on Per- idermium harknessii aecia. Mountainous loca- tions and hosts from which the specimens were collected indicate they all could be from pine stem rust aecia. Specimens of Phalacropsis dispar collected during this study have been deposited in the Museum of Comparative Zoology (Agassiz Museum), Harvard University, and the National Mu- seum of Natural History Smithsonian In- stitution, Washington, D.C. Distribution The distribution of P. dispar observed in this study is indicated in Figure 3 and Table 1. With few exceptions, the beetle was found wherever pine stem rust fungi were examined closely during the aecial sporulation period. The most intensive study was made in Utah and in the Sierra Nevada of California, and the number of sites for the beetle in these areas (Fig. 3) is a reflection of this and is not necessarily an indication of abundance. During a five-year study of western gall rust on pines in coastal areas of California, P. dispar was not encountered, nor was it listed by Byler et al. (1972a), who studied the same rust in these areas. In north coastal California near Fort Bragg, however, an unidentified larva of similar habit was abundant in aecia of Peridennium harknessii on Pinus contorta Fig. 3. Distribution of Phalacropsis dispar observed in this study on pine stem rust fungi in the western United States. 374 Great Basin Naturalist Vol. 42, No. 3 ssp. bolanderi (Pari.). Several attempts to rear adults failed. In the Sierra Nevada, Phala- cropsis dispar was found on Peridertniiim fila- mentoswn Inyo form (Peterson 1968), P. sta- lactiforme both forms, P. harknessii, Cronartium occidentale, and "Bethel" blister rust (Dixon 1978) (Table 1). Cronartium com- andrae was not studied in this area. In the Bucks Lake area, Pluikicropsis dispar was not found in aecia of the introduced C. ribicola on Piniis Imnbertiana (Dougl.). In this area, the C. ribicoki -infected P. lambertiana oc- curred in several stands that were intermixed with Peridermium harknessii-iniected Piniis ponderosa and P. contorta. The aecia of Per- idennium Iwrknessii on these pines were in- fested with Phalacropsis dispar. In the Spring Moimtains of southern Nevada, P. dispar was found on the Coronado and Powell (Peterson 1968) forms of Peridermium filatnentosum limb rust and was especially abimdant on the albino form (Mielke and Peterson 1967) of western gall rust on Pinus ponderosa. In Ari- zona, Phalacropsis dispar was observed on the Coronado and Powell forms of Per- idermium filamentosum limb rust on Pinus ponderosa on the Kaibab Plateau and in the San Francisco Mountains. In Utah, Phala- cropsis dispar occurred on the Coronado and Powell forms of Peridermium filamentosum limb rust on Pinus ponderosa in the Abajo Mountains and on the Aquarius and Mark- agunt Plateaus in the southern end of the state. It also occurred on the Powell Per- idermium filamentosum limb rust on Pinus ponderosa at the eastern end of the Uinta Mountains in northeastern Utah. It was not found on C. comandrae, Peridermium hark- nessii, and P. stalactiforme on Pinus contorta in the Wasatch Mountains of northern Utah. Phalacropsis dispar was found— although rarely— on C. comandrae on Pinus contorta on the Cassia Plateau of southern Idaho and on the northern end of the Wind River Range in western Wyoming. It occurred in Per- idermiuni harknessii aecia on Pinus contorta in the Island Park area of eastern Idaho and on the same rust on both P. contorta and P. ponderosa in the Sawtooth Mountains in the south central part of the state. It was found Table 1. The Peridermium species of pine stem nist fungi found in this study to be infested by Phalacropsis dispar in the western United States. Peridermium rust Pine host Location Cronartium coleosporioides Arth. Periderrnimn filamentosum Peck Coronado hnib rust Powell limb rust Inyo limb rust Peridermium stalactiforme Arthur & Kern Stalactiforme limb rust Stalactiforme canker rust Endocronartium harknessii (J. P. Moore) Y. Hiratsuka {Peridermium harknessii J. P. Moore) Western gall rust Albino western gall rust Cronartiitm comandrae Peck Peridermium pyriforme Peck Comandra blister rust "Bethel" blister rust Cronartium occidentale Hedg., Bethel, & Hunt Peridermium occidentale Hedg., Bethel, & Hunt Pinyon blister rust Pinus ponderosa Pinus ponderosa Pinus jeffreyi Pinus jeffreyi Pinus contorta Pinus ponderosa Pinus contorta Pinus ponderosa Pinus ponderosa Pinus contorta Pinus contorta Pinus monophylla Arizona, Nevada, Utah Arizona, Nevada, Utah California California, Nevada California, Oregon Idaho California, Idaho, Oregon California, Idaho Nevada, Utah Idaho, Wyoming California California September 1982 Nelson: Natural Control of Pine Stem Rust 375 on a single canker of Peridermium stalacti- forme on Pinus ponderosa in the Salmon Riv- er Mountains. In the Malheur National For- est of Oregon, Phalacrapsis dispar was found on Peridermium harknessii and P. stalacti- forme on Pinus contorta. In the central Idaho and Oregon areas mentioned, larvae and adults of an Epuraea sp. (another aeciospore- consuming beetle) were also present. From northern Utah northward, occurrence of Phalacropsis dispar was less frequent to rare and southward from this point it was increas- ingly abundant. Life History Observations Phalacropsis disbar was first noted in mid- May near Bucks Lake and in late May at Lee Vining and Red Canyon. Aecia of the associ- ated rust fungi were just beginning to appear through the bark. On simny days, adult bee- tles were seen on rusted limbs, crawling about and copulating. Egg-laying by the beetle was observed only in association with Peridermium filamentosum on Pinus jeffreyi in the Sierra Nevada of California. Single eggs were deposited at the exterior base of aecia. Larval activity began before full devel- opment of aecia and well in advance of per- idial rupture. With the beginning of aecio- spore release, larval activity had reached a peak. Beyond this stage, in the case of early sporulating rust fungi, adult beetles were not seen on rusted limbs; however, in the case of later sporulating Powell and Inyo forms of Peridermium filamentosum, adults were seen throughout July and August well after aecial maturity. After larval maturity, the insect disappeared from rusted limbs and galls and possibly pupated in the needle duff at the base of trees. Whether or not the insect over- winters in the pupal or adult stage in its nat- ural environment was not studied. In laboratory tests (Table 2), rusted stems were collected 10 to 15 days after adults were first seen. The first larvae began mov- ing from aecia about 6 days after rust speci- mens were placed in glass beakers in the lab- oratory. Pupation began 4 to 7 days later on the floor of the beaker under masses of fecal debris, unconsumed aeciospores, and bark fragments that dropped from rusted stems. Beginning of pupation was determined by a shortening and thickening of larvae, their im- mobility, and the appearance of black eye spots. The first adults emerged 14 to 36 (av- erage 23) days after the first larvae began leaving aecia. If development proceeds sim- ilarly under natural conditions, roughly 30 to 40 days would be required for completion of the beetle's life cycle. The beetles were not fouind on other plants in the vicinity of the pine stem rusts, al- though members of the family Phalacridae are known to frequent flowers of the Com- positae in particular (Amett 1973). Beetle Infestations and Damage At the Bucks Lake location where P. hark- nessii occurred on Pinus ponderosa, several hundred galls were observed annually on a stand of young trees covering approximately 0.5 ha. During the entire study period, not a Table 2. Incubation period of Phalacropsis dispar in laboratory tests. Larvae Total days Rust' Placed moving from first fungus Collection in from Pupation First larval Year National forest pine host date beaker aecia began adults maturity 1967 Toiyabe, California Ps/Pj 7/14 7/17 7/20 7/24 8/3 14 1968 Toiyabe, California Ps/P] 6/22 6/25 7/1 7/5 7/17 16 1969 Dixie, Utah Pf/Pp 6/23 6/25 7/3 7/22 7/28 25 1970 Plumas, California Ph/Pp 5/26 5/28 6/3 6/18 6/29 26 1971 Plumas, California Ph/Pp 5/30 6/6 6/12 6/16 7/14 36 1973 Plumas, California Ph/Pp 5/28 5/28 6/4 6/11 6/26 22 1974 Plumas, California Ph/Pp 5/23 5/25 5/31 6/4 6/18 18 1975 Toiyabe, Nevada Pha/Pp 7/12 7/13 7/18 7/25 8/6 19 Ps/ Pj = Peridermium stalactiforme/ Pinus jeffreyi Pf/Pp = Peridermium filamentosum /Pinus ponderosa Ph/Pp = Peridermium harknessii/ Pinus ponderosa Pha/Pp = Peridermium harknessii (albino) /Pinui ponderosa 376 Great Basin Naturalist Vol. 42, No. 3 single gall was found without some evidence of beetle larval activity. Larvae reduced ae- ciospores and the underlying sporogenous mycelium to a mass of fecal debris (Fig. 2B) within approximately two weeks. The initial sporulation of young galls is usually a week or so later than older galls; these also become infested. In a single laboratory test (1971), four galls, 4 to 8 cm in diameter, produced an average of 130 adult beetles per gall. A beetle infestation of similar intensity oc- curred on Peridermium stalactiforme on Finns jeffreyi at Lee Vining Creek. Aecia matured about mid-June and were com- pletely destroyed within about two weeks. Along a three km stretch of Lee Vining Creek, not one of the 20 limb-rusted trees ex- amined annually remained free of the beetle. Aecial confluency with this rust is markedly less than with western gall rust, but few aecia were uninfested. Those that were uninfested were the more scattered aecia among needles toward the distal end of limbs. In two labora- tory tests (1968 and 1971), a total of 3.9 m of typical msted limbs yielded 3.4 adult bee- tles/cm and a total of 4.6 m of rusted limbs yielded 1.2 beetles/ cm. Beetle infestation of the Powell Per- idermium filamentosum studied at Red Can- yon was somewhat different from the in- festations of P. harknessii and P. stalactiforme described above. Vv^ith this rust fungus, aecia are tongue shaped and single with little con- fluency (compare Figs. 2 A and 2C). Seldom did more than one beetle larva occur within a single aecium. Larvae consumed the basal mycelium of an aecium first and then moved into the upper part. Frequently the upper part of aecia remained untouched. Pressure from the growing fecal mass often burst the aecium at its base. It appeared that a single aecial mass was sufficient to rear a single lar- va. Usually, fewer than 50 percent of the aecia became infected on rusted limbs of the 12 trees observed annually. The amount of damage seemed to vary more from year to year with this rust than with P. harknessii and P. stahctiforme. In California, few Inyo P. filamentosum aecia were infested some years, but virtually all aecia became infested in other years. The other pine stem rusts were studied less. Infestation of P. stalactiforme on Pinus contorta in the Sierra Nevada of California was as severe as that noted on P. jeffreyi at Lee Vining Creek. Infestation of the Coro- nado Peridermium filamentosum was similar to that of the other forms of P. filamentosum, although limited to a shorter sporulation pe- riod. Cronartium comandrae was studied little, and so the extent of infestation was not known. Observation of the beetle on C. occi- dentale on Pinus monophylla Torr. and Frem. was limited to a large outbreak of the rust near Monitor Pass, Alpine County, in the Sierra Nevada of California. During the three seasons observed, aecial spore masses were virtually destroyed by the beetle in all trees examined. Discussion The beetle Phalacropsis dispar is reported here to consume the aeciospores and the ae- cial mycelium of all species of native western pine stem rust fungi except Cronartium con- igenum (Pat.) Peterson and C comptoniae, neither of which was investigated. Aecial fructifications not only provide food but also protective chambers for the larvae. Within several weeks' time, aecia may be completely ravaged of their content; only the peridial shell and masses of fecal debris remain. Based on laboratory tests and field observations, the life cycle of the beetle is completed in 30 to 40 days. Apparently, the beetle overwinters as an adult and emerges in the spring to mate and lay eggs as aecia. of the rust fungi begin forming. The rarity of P. dispar in insect col- lections is further evidence it is highly specif- ic to these rust fungi. The insects are abun- dant on pine stem rust fungi, but these fungi are rather obscure to insect collectors. Also, if they live on other fungi or the flowers of vascular plants as do other Phalacaridae, they probably would turn up more frequently in collections. The distribution of P. dispar appears to be primarily south of the 40th parallel. North of the parallel, the niche becomes increasingly occupied by Epuraea spp. Some adults or lar- vae of P. dispar could be found in almost all areas where native pine stem rust fungi in the aecial sporulation state were studied. The beetle appeared, however, to be absent in some areas such as coastal California and September 1982 Nelson: Natural Control of Pine Stem Rust 377 northern Utah. R. S. Peterson (pers. comm.) has found Phalacropsis to be common to abimdant on both the albino and orange- spored Peridermium harknessii in the Black Hills of South Dakota and Wyoming and in parts of Colorado. Reduction of the aeciospore inoculum by Phalacropsis dispar was not determined in quantitative terms, but based on my observa- tions, it can be a high percentage of the po- tential amount. Not only is there a reduction of spores already formed at the time larvae begin feeding, but consumption of the under- lying sporogenous mycelium reduces the final quantity of spores produced as well as the length of the sporulation period. Quantitative field data are needed to establish what im- pact P. dispar has in reducing aeciospore in- oculum and in the natural control of pine stem rust fungi. Nevertheless, all factors in the host-pathogen-environment interaction are of some importance. Information on the aeciospore dispersal period for native western pine stem rust fungi is limited. R. S. Peterson (1959), study- ing spore release from Peridermium Imrk- nessii at high elevation sites in Colorado, found intermittent, but abundant, sporulation for one to two months. In a two-year study, G. W. Peterson (1973) trapped 88 and 91 percent of the season's total release of P. harknessii aeciospores during a two- and three-week period each May. The remainder (9 to 12 percent) was trapped through June. In Krebills's (1968) study of C. comandrae in the Rocky Mountain States, aeciospore dis- persal began in late May, peaked in the last half of June, and usually continued in small amounts through summer. With increasing elevation, dispersal was delayed and the peri- od shortened. The tremendous mass of spores produced by the aecial state of the pine stem rust fungi is an evolutionary adaptation that increases the chance for survival of these propagules through the environmental hazards of dis- semination. Aeciospores (annual inoculum) serve as the initial step in what is a long series of events leading to infection of pine. They also function in the spread of rust fungi over long distances, and a reduction in the in- oculum load would seem to be important. In autoecious rust fimgi, such as Endocronar- tium harknessii (Peridermium harknessii), re- duction of the aecial inoculum is likely to be of more significance than it would be in het- eroecious rust fungi, such as C. coleospo- rioides or C. comandrae, which have the ure- dinial multiplication phase. The Inyo and Powell forms of P. filamentosum, with aecia adapted to survive potentially long dry peri- ods, could be severely affected by early de- struction of aecia. Even with heteroecious rust fungi, in the more arid regions of the western United States where favorable mois- ture requirements for infection are likely to occur less frequently during the uredinial phase, intensity of infection by aeciospores would seem to be important. Phalacropsis dispar is possibly specific to native conifer stem rust fungi, including C. occidentale, which is similar in host range to C. ribicola. If P. dispar does not feed on the introduced C. ribicola, it may reveal a deficit in the natural control of this rust fungus in North America. From reports in the literature, there appears to be a larger diversity of secondary or- ganisms that inhabit native western pine stem rusts (Wollenweber 1934, Powell 1971a,b,c,d,e, Byler et al. 1972b, Powell et al. 1972, Powell 1974, Hiratsuka et al. 1979, Tsuneda and Hiratsuka 1979, 1980) than in- habit C. ribicola blister rust (Mielke 1933, 1935, Kimmey 1969, Williams 1972, Furniss et al. 1972, Hungerford 1977, Hunt 1978). Members of the genus Epuraea evidently are widely distributed in the northwestern United States and western Canada (Hatch 1961, Parsons 1967, Powell 1971d, Powell et al. 1972). The aggressive Eptiraea obliquus and E. ovata, obligate consumers of aecio- spores of four different species of native pine stem rust fungi (Hubert 1923, Powell et al. 1972), were not reported by Furniss et al. (1972) to occur on C. ribicola blister rust can- kers in northern Idaho. Both E. obliquus and E. ovata are known to exist in the eastern Oregon, northern Idaho, and western Mon- tana areas (Hubert 1923, Hatch 1961). The absence of natural control of C. ribicola in western North America has been attributed primarily to a lack of host resistance (Bing- ham et al. 1971, Hoff and McDonald 1972). Byler et al. (1972a) provide evidence that secondary fungi and insects were primarily 378 Great Basin Naturalist Vol. 42, No. 3 responsible for stabilizing populations of Per- idermium harknessii at a low equilibrium po- sition on coastal California pines. Pinus radi- ata and other coastal pine species become heavily infected with this rust fungus. Al- though these species are apparently highly susceptible to infection, resistance mecha- nisms probably provide tolerance (True 1938). Genetic drift of susceptibility from areas protected by an environment unfavor- able for infection could account for part of this high susceptibility. The extent of natural control resulting from the activity of second- ary organisms is a possible explanation for the reduced need for host resistance. The rust fungi are well-known obligate parasites and have evolved with their hosts to a state of mutual survival. Considering the factors involved in rust disease epidemiology, it appears that secondary organisms would tend to reduce disease incidence and thus re- duce selection for host resistance. The impor- tance of secondary organisms should perhaps be placed more in the perspective of what disease incidence might be in their absence rather than as candidates for biological con- trol above and beyond what occurs in a natu- ral environment. Literature Cited Anderson, N. A., and D. W. French. 1964. Debarking jack pine infected with sweetfem rust. Plant Dis. Kept. 48:530-531. Arnett, R. 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A Peridennium rust associated with dwarf mistletoe on lodgepole pine. Unpublished thesis. Colorado State Univ. 100 pp. FuRNiss, M. M., R. D. Hungerford, and E. F. Wicker. 1972. Insects and mites associated with western white pine blister rust cankers in Idaho. Cana- dian Entomol. 104:1713-1715. Hanover, J. W. 1966. Tree improvement for disease re- sistance in western United States and Canada. Pages 53-66 in Breeding pest resistant trees. Proc. NATO-IUFRO Sym., Penn. St. Univ. 1964. Pergamon Press, Oxford, England. 505 pp. Hatch, M. H. 1961. The beetles of the Pacific North- west. Part III: Pselaphidae and Diversicornia 1. Univ. of Washington Press, Seattle. 503 pp. Hiratsuka, Y., and J. M. Powell. 1976. Pine stem rusts of Canada. Canada Dep. Environ., For. Serv., Forestry Tech. Rep. 4. 83 pp. Hiratsuka, Y., A. Tsuneda, and L. Sigler. 1979. Occur- rence of Scytalidium uredinicola on Endocronar- tiiim harknessii in Alberta, Canada. Plant. Dis. Rept. 63:512-513. Hoff, J. R., and G. I. McDonald. 1972. Stem rusts of conifers and the balance of nature. Pages 525-536 in Biology of rust resistance in forest trees. Proc. NATO-IUFRO Adv. Study Inst., Moscow, Idaho, 1969. USDA For. Serv. Misc. Publ. 1221. 681 pp. Hubert, E. E. 1923. The life history of Cronartium co- leosporioides. Unpublished manuscript. Div. For. Dis. Res., Washington Off. For. Serv. 76 pp. Hungerford, R. D. 1977. Natural inactivation of blister rust cankers on western white pine. For. Sci. 23:343-350. Hunt, R. S. 1978. Slugs feeding on Cronartium in British Columbia. Canadian For. Serv. Res. Note 34:21. Kimmey, J. W. 1969. Inactivation of lethal-type blister rust cankers on western white pine. J. For. 67:296-299. Krebill, R. G. 1968. Cronartium comandrae in the Rocky Mountain States. USDA For. Serv., In- termt. For. and Range Expt. Sta., Res. Pap. INT- 40, 28 pp. KuHLMAN, E. G. 1981a. Mycoparasite effects of Scytali- dium uredinicola on aeciospore production and germination of Cronartium quercuum sp. fusi- fonne. Phytopathology 71:186-188. 1981b. Parasite interaction with sporulation by Cronartium quercuum subsp. fusiforme on lob- lolly and slash pine. Phytopathology 71:348-350. KuHLMAN, E. G., J. W. Carmichael, and T. Miller. 1976. Scytalidium uredinicola a new mycopara- site of Cronartium fusiforme on Pinus. Mycologia 68:1188-1194. KuHLMAN, E. G., AND T. MiLLER. 1976. Occurrence of Tuberculina maxima on fusiform rust galls in the southeastern United States. Plant Dis. Rept. 60:627-629. Leaphart, C. D., AND E. F. Wicker. 1968. The in- effectiveness of cycloheximide and phytoactin as chemical controls of the blister rust disease. Plant Dis. Rept. 52:6-10. LeConte, T. L. 1879. The Coleoptera of the alpine Rocky Mountain Regions— Part II. Bull. U.S. Geol. Survey 5(3,XXV):499-513. September 1982 Nelson: Natural Control of Pine Stem Rust 379 Merkel, E. p. 1958. Diortjctrio cone moth attack as re- lated to cone nist of slash pine in North Florida. J. For. 56:651. MiELKE, J. L. 1933. Tuherculina maxima in western North America. Phytopathology 23:299-305. 1935. Rodents as a factor in reducing aecial spo- rulation of Cronartium rihicola. J. For. 33:994-1003. MiELKE, J. L., AND R. S. Peterson. 1967. Albino Per- idennium harknessii in ponderosa pine. Plant Dis. Rept. 51:306-509. Myren, D. T. 1964. Insects and fimgi associated with Cronartium fusiforme-iniected tissue and com- parisons of the strength of infected and healthy wood. Phytopathology 54:902. Nighswander, J. E., AND R. F. Patton. 1965. The epidemiology of the jack pine-oak gall rust {Cro- nartium quercuum) in Wisconsin. Canadian J. Hot. 43:1561-1581. Parsons, C. T. 1967. North American Nitidulidae (Co- leoptera) IV. Epuraea associated with hingi on pine and oak. Canadian Entomol. 99:734-737. Peterson, G. W. 1973. Dispersal of aeciospores of Per- idennium harknessii in central Nebraska. Phyto- pathology 63:170-172. Peterson, R. S. 1959. Pine gall mst in the Rocky Moun- tains. Dissertation. Univ. of Michigan. Diss. Abstr. 20(5)1557, 111 pp. 1966. Limb rust damage to pine. USDA For. Serv., Intermt. For. and Range Expt. Sta., Res. Pap. INT-31, 10 pp. 1968. Limb rust of pine: the causal fimgi. Phyto- pathology 58:309-315. Peterson, R. S., and F. F. Jewell. 1968. Status of American stem rusts of pine. Ann. Rev. Phyto- pathology 6:2.3-40. Powell, J. M. 1971a. Fungi and bacteria associated with Cronartium comandrae on lodgepole pine in Alberta. Phytoprotection 52:45-51. 1971b. Occurrence of Tuberculina maxima on pine stem rusts in western Canada. Canadian Plant Dis. Surv. 51:83-85. 1971c. Incidence and effect of Tubercu- lina maxima on cankers of the pine stem rust, Cronartium comandrae. Phytoprotection 52:104-111. 1971d. The arthropod fauna collected from the comandra blister mst, Cronartiinn comandrae on lodgepole pine in Alberta. Canadian Entomol. 103:908-918. 1971e. Additional records of Mycodiplosis larvae (Diptera; Cecidomylidae) feeding on rust fungi. Canadian Plant Dis. Surv. 51:86-87. 1974. The role of natural biological agents in con- trolling a pine stem rust {Cronartium comandrae). Rlue Jay 32:75-79. Powell, J. M., H. R. Wong, and J. C. E. Melvin. 1972. Arthropods collected from stem ru.st cankers of hard pines in western Canada. Canada Dept. En- viron., Canadian For. Serv., Northern For. Res. Centre, Edmonton, Alberta. Rep. NOR-X-42, 19 pp. Snell, W. H. 1919. Observations on the relation of in- sects to the dissemination of Cronartium rihicola. Phytopathology 9:451-464. Toko, H. V., D. A. Graham, C. E. Carlson, and D. E. Ketcham. 1967. Effect of past Ribes eradications on controlling white pine blister rust in northern Idaho. Phytopathology 57:1010. True, R. P. 1938. Gall development on Pinus sylvestris attacked by the Woodgate Peridermium, and morphology of the parasite. Phytopathology 28:24-49. Tsuneda, a., and Y. Hiratsuka. 1979. Mode of para- sitism of mycoparasite, Cladosporium galUcoIa, on western gall rust, Endocronartium harknessii. Canadian J. Plant Pathol. 1:31-36. 1980. Parasitization of pine stem rust fungi by Monocillium nordinii. Phytopathology 70:1101-1103. Tsuneda, A., Y. Hiratsuka, and P. T. Maruyama. 1980. Hyperparasitism of Scytalidium uredinicoki on western gall rust, Endocronartium harknessii. Ca- nadian J. Hot. 58:1154-1159. Williams, R. E. 1972. Fungi associated with blister rust cankers on western white pine. Dissertation. Washington State Univ. 125 pp. Xerox Univ. Mi- crofilms, Ann Arbor, Mich., No. 72-31308. Diss. Abstr. 33/05-B, p. 2434. WoLLENWEBER, H. W. 1934. Fusorium bactridioides sp. not), associated with Cronartium. Science 79(2060):572. Van Arsdel, E. P. 1961. Growing white pine in the Lake States to avoid blister rust. USDA For. Serv. Lake States For. Exp. Sta., Pap. 92, 11 pp. Wong, H. R. 1972. Dioryctria banksiella (Lepidoptera: Pyralidae) in the western gall rust, Endocronar- tium harknessii (Basidiomycetes: Uredinales). Ca- nadian Entomol. 104:251-255. SPECIES-HABITAT RELATIONSHIPS IN AN OREGON COLD DESERT LIZARD COMMUNITY David F. Werschkul' Abstract.- The abundance and diversity of lizards in nine habitat types from Oregon were studied from May through October 1980. Eight species were from eight habitat types. The most common species were Sceloporous oc- cidentalism Uta stanshuriana, Scelopowm graciosus, and Cnemidophorus tigris. Phrifnosoma douglassi was uncommon and Eiimeces skiltonianus was not observed. Temporary streams in nonbasaltic areas were the most productive habi- tat m terms of Hzard abundance but sagebrush areas were the most productive habitat in terms of species diversity No hzards were recorded from grassland conversion areas. The conflict between a land management policy that em- phasizes both vegetation conversion and conservation of present wildlife stocks is discussed. The herpetofauna of Malheur County, lo- cated in the extreme southeastern corner of Oregon, has been largely neglected in spite of the biological interest in this transition zone between the Great Basin and the cold desert areas of northern Oregon, Washington, and Idaho (Storm and Pimentel 1949, Fergu- son et al. 1958, St. John 1980). The purpose of this study was to survey the herpetofauna and relate abundance and diversity to habitat type. Herein I report my findings on the structure of the lizard community. The abun- dance and diversity of amphibians and snakes is reported elsewhere (Werschkul 1980). Study Area The climate of southeastern Oregon is characterized by variation with short, hot summers, mean July temperature of 25 C, and long, cold winters, mean temperature be- tween November and March of 4.6 C (Loy et al. 1976). Rainfall is highest in May, 40 mm, and lowest in August, 6 mm. A complex geo- logic area, early Miocene extrusions of basalt and rhyolite form the foundations for present formations (Kittleman et al. 1967, Kittleman 1973). Bisected by erosion, these volcanic platforms have formed elongate ridges and basins with deposition of alluvial materials along river channels. Plant communities from this high desert ecosystem, the juniper-sage- brush woodland extension of the Great Basin pinyon-juniper woodland (Detling 1968), have been classified by Franklin and Dymess (1973), and, although big sagebrush (Arte- misia fridentata) /hluehunch wheatgrass (Agropyron spiratum) is recognized as the cli- matic climax (Eckert 1957, Tueller 1962), na- tive grasses have largely disappeared because of livestock grazing (Tueller 1962). Regard- less, the composition of the plant community responds to local conditions and black sage- brush [Artemisia arhiiscula) is found on shal- low soils, shadscale (Atriplex confertifolia) and hopsage (Grayia spinosa) are common on xeric sites, and greasewood (Sacrobatiis ver- miculatus) dominates on sandy and alkaline sites. Methods The areas censused were bordered to the east by the Oregon-Idaho state line (117°05'W), to the west by the Alvord Desert (118°30'W), to the south by the Oregon- Nevada state line (42°00'N), and to the north by the Malheur River (44°00'N) from May through October 1980 (Fig. 1). Each census site was classified into one of nine habitat types depending on the composition of the plant community and the geologic and soil conditions. These habitat types were sage- brush, alkaline flats, grasslands, rocky areas of basaltic or nonbasaltic origin, temporary streams in basaltic or nonbasaltic areas, per- manent streams, and sand dunes. Sagebrush.— A composite category for sagebrush areas including higher elevation sites of sagebrush in association with bitter- 'Kalmiopsis Field Station, Agness, Oregon 97406. 380 September 1982 Werschkul: Cold Desert Lizard Community 381 brush {Purshia tridentata), juniper {Juniperus occidentalis), and mountain mahogany {Cer- cocarpus ledifolius); xeric sites with saltbrush {Atriplex sp.); and rocky areas with black sagebrush, as well as the more common plateau and basin regions with nearly pure stands of big sagebrush. Alkaline flats.— The shrub community from these ancient lakebeds, or playas, in- clude shadscale, hopsage, and greasewood, al- though on extremely alkaline areas the vege- tation was absent except on small (ca 10 m^) hummocks. Grasslands.— Native grasses have largely disappeared because of livestock grazing. Range restoration projects, under the admin- istration of the Vale District of the Bureau of Land Management, include seeding of crested wheatgrass {Agropyron cristatum) (Heady and Bartolome 1977), and it is these areas I censused during this study. Rocky areas of basaltic origin.— Those rocky areas that included exposed bluffs, talus slopes of basaltic rubble, and the more recent basaltic extrusions of the smooth pa- hoehoe type. Rocky areas of nonbasaltic origin.— A composite category that included areas as di- verse as arkose sandstone and hard pyroclas- NEVADA Fig. L Lx)cation of survey, Malheur County, Oregon. tic flows. Shrub cover was always sparse, less than one shrub stem per 25 m^. Sand dunes.— Sandy areas devoid of vege- tation except for border cover, usually greasewood or big sagebrush. Temporary streams in basaltic areas.— Streams on ridges and steep slopes usually eroded to basalt; sandy areas might be pres- ent, but they were not extensive. The most common woody plants were willows {Salix spp.) and chokecherry (Primus sp.). Temporary streams in nonbasaltic AREAS.— Streams on relatively flat areas usu- ally did not erode to basalt, although basaltic boulders were sometimes present. Woody plants were for the most part absent and the plant community was similar to that found in sandy, slightly alkaline areas and included greasewood, rabbitbrush {Chrysothamnus spp.), and Indian ricegrass (Oryzopsis hymenoides). Permanent streams.— Those areas adja- cent to permanent flowing water. Usually, a well-developed riparian community of wil- lows, Cottonwood {Populus trichocarpa), and hawthorn {Crataegus spp.) was present. Lizards were censused by slowly walking (ca 0.5 m sec-i) through areas noting those species present. Each census period lasted 20 minutes, and animal abundance was calcu- lated as the number of animals seen per cen- sus period. In general, animals were not pur- sued or captured unless identification was uncertain. An area might be censused repeat- edly, but no particular site was censused more than once. For example, Jordan Cra- ters, a large expanse of recent basaltic extru- sions (Kindschy and Maser 1978), was cen- sused for 17 census periods, although no particular site was visited more than once. Most censuses were made during the morn- ing, 0630-1030, and the afternoon, 1500-1900, although some censusing oc- curred at other times. Malheur County, the study area, is large, approximately 2.6 million ha. Consequently, areas censused were representative of the habitats found there. Some areas were chosen because of reports of one species or another. To minimize error in determining species- habitat relationships, I censused each habitat type often enough, a total of 358 census peri- ods or 4160 minutes, to reduce the impacts of 382 Great Basin Naturalist Vol. 42, No. 3 season, time of day, and weather on animal activity (see Mayhew 1968, Parker and Pian- ka 1976). It is recognized that the Phryno- soma are probably underestimated and Eu- meces may have been missed due to the survey technique used here. Results and Discussion Nine species comprise the lizard fauna of southeastern Oregon (Stebbins 1954, St. John 1980), eight of which were recorded during this study (Table 1). The most commonly ob- served species were Sceloporous occidentalis, Uta stansburiana, Sceloporous graciosus, and Cnemidophorus tigris. Phyrnosoma douglassi were uncommon, only two sightings, and Eu- meces skiltonianus were not observed (but see St. John 1980). A total of 192 lizards or 0.5 animals per census period was recorded. Habitat importance values were calculated in two ways: (1) as the percent of census sites per habitat type with at least one animal sighted, and (2) the average number of ani- mals per successful census site (those sites with at least one animal sighting). As might be expected, these methods of estimating spe- cies-habitat values are positively correlated (Spearman Rank Correlation = 0.48; p < 0.01; n = 29), although some noteworthy ex- ceptions exist. For example, for temporary streams in nonbasaltic areas S. occidentalis were found in 9.1 percent of the census sites, a relatively low value, but averaged 2.7 ani- mals per successful census site. This may have resulted from the fact that, when large boulders (used for sunbathing) were present, S. occidentalis were more commonly ob- served than when large boulders were absent. Likewise, S. graciosus were relatively uncom- mon in sagebrush areas, 4.4 percent of the census sites, although relatively high numbers were found when encountered, 2.1 animals per successful census site. In this case, S. gra- ciosus were uncommonly observed except where small mammal burrows were present. These lizards apparently use these burrows to aid in thermoregulation and to avoid pred- ators. Conversely, although Gambelia wisli- zeni were commonly observed from tempo- rary streams in nonbasaltic areas, 18.1 percent of census sites, numbers were always low, 1.5 animals per successful census site. Gambelia wislizeni, however, were not Table 1. Occurrence of lizards from nine habitat types from Malheur County, Oregon. Habitat -Q ;s Species Crotaphytits bicinctores Gambelia wislizeni Sceloporus occidentalis Sceloporus graciosus Uta stansburiana Phnjnosorna douglassi Phnjnosoma platijrhinos Cnemidophorus tigris Nb 10 20 1.3c 2.00d - 18 25 5.0 1.28 11.1 1.50 20 40 2.5 1.23 - 16 32 4.4 2.10 33.3 2.30 16 38 1.9 1.70 - 2 2 1.3 1.00 - 8 10 2.5 1.00 11.1 1.50 19 30 5.0 1.70 5.5 1.00 fNumber of census periods with at least one animal sighting, ^otal number of animals sighted. percent of census sites with at least one animal sighting. Average number of animals per successful census period. o o >. -a c C/5 ^1 r- CD H .S o -Q u a. 7.1 15.1 1.00 12.5 2.60 18.1 2.00 1.50 19.5 7.1 7.7 9.1 6.4 2.33 1.00 1.00 2.67 2.00 6.8 1.00 33.3 6.4 2.70 1.50 2.2 1.00 12.5 1.50 25.0 2.00 12.1 1.30 3.2 1.00 September 1982 Werschkul: Cold Desert Lizard Community 383 observed in high numbers in any habitat (Table 1) and it is suspected that some spac- ing mechanism may be at work. Overall, when microhabitat characteristics cause clumping, such as the presence of boulders for S. occidentalis or small mammal burrows for S. graciosus, then the number of animals per successful census site tend to over- estimate habitat importance. Sagebrush habitat, in terms of species ob- served, was the most productive census area, with all eight species observed (Table 1). Grasslands, with no observations, was the least productive. Shrub cover is important in determining lizard distribution and abun- dance patterns by providing shaded areas needed for thermoregulation and as hiding sites from predators (Germano and Hun- gerford 1981). Shrub cover removal, by re- ducing the vertical and horizontal habitat stratification and shading and hiding areas, results in poor habitat for lizards. In terms of abundance, the habitat associated with tem- porary streams in nonbasaltic areas was the most productive and grasslands the least pro- ductive (Table 2). Each species had a pre- ferred habitat (Table 1), and that associated with temporary streams in nonbasaltic areas was preferred by more species, three, than any other habitat type. Although sagebrush areas had the highest number of species ob- served, only P. douglassi preferred this habi- tat type (Table 1) and the average number of lizards seen per census period was com- paratively low (Table 2). The complete lack of observations of liz- ards in grasslands was unexpected. Although studies of lizard-habitat relationships in Ari- zona have shown low use of grasslands areas by most species, some species (e.g., Sonora spotted whiptail, Cnemidophorus sonorae) were more common in grasslands than else- where, and only a minority of species (4 of 9) went unrecorded in grasslands (Germano and Hungerford 1981). Two possible factors con- tributing to the lack of lizard sightings in the grasslands from this study were: (1) I ex eluded, as much as possible, any edge effects during censusing by picking grassland areas away from other habitat types and areas lacking sagebrush, boulders, or any other fea- tures causing habitat stratification, and (2), during the study year, rainfall was high, caus- ing dense and tall stands of grasses to devel- op. This increased the technical difficulties associated with the walking transect survey and may have caused local lizard movements to habitats with open areas needed for forag- ing. It seems unlikely that all species of liz- ards avoided grasslands at all times, and some species may benefit from increases in the acreage of grasslands (Germano and Him- gerford 1981). Census of areas with habitat discontinuities may have shown more use of grasslands by lizards than indicated here. Still, to a large extent, lizards failed to use grasslands in this study area and a question arises relative to the economic necessity for expanding grasslands and the desire to con- serve wildlife stocks. Grasslands, important for livestock production, were avoided by liz- ards in this study, and other plant commu- nities, important to lizards, are unproductive for livestock. My findings suggest that, when Table 2. The abundance and diversity of lizards for nine habitat types from Malheur County, Oregon. Diversity (H) is measured as the Shannon-Weaver index (H = 2 -pj In pj). Habitat Temporary streams in nonbasaltic areas Alkaline flats Sand dunes Rocky basaltic areas Sagebrush Permanent streams Temporary streams in basaltic areas Rocky nonbasaltic areas Grasslands Number of census periods Number of species Average number of lizards per census period 33 5 1.41 1.97 18 4 0.97 1.17 16 3 1.01 0.94 46 4 0.83 0.61 156 8 1.87 0.36 31 3 0.94 0.26 13 1 _ 0.08 14 1 — 0.07 18 0 - 0.00 384 Great Basin Naturalist Vol. 42, No. 3 multiple use management is desired, con- version areas need to maintain some charac- teristics of the native vegetation. For ex- ample, areas adjacent to temporary streams, frequently not suitable for crested wheat- grass, if conserved, would reduce the costs to the lizard community resulting from vegeta- tion control programs. Importantly, dispersal corridors for the lizards, as well as valuable habitat for other wildlife, would be con- served. Finally, sagebrush areas are especially important to lizards, because all species were found to use this habitat. Management pro- grams should follow the advice of Germano and Hungerford (1981) and clear variously shaped areas rather than attempt total re- moval of all nongrass vegetation. Acknowledgments Bob Kindschy provided information on hz- ard locations in southeastern Oregon. This study was supported by the Bureau of Land Management (OR 910-CTO-14). Literature Cited Detling, L. E. 1968. Historical background of the flora of the Pacific Northwest. Univ. of Oregon Mu- seum of Natural History. Eugene. 57 pp. EcKERT, R. E., Jr. 1957. Vegetation-soil relationships in some Artemisia types in northern Harney and Lake counties, Oregon. Unpublished dissertation. Oregon State College, Corvallis. Ferguson, D. E., K. E. Payne, and R. M. Storm. 1958. Notes on the herpetofauna of Baker County, Ore- gon. Creat Basin Nat. 18:63-65. Franklin, J. F., and C. T. Dyrness. 1973. Natural vege- tation of Oregon and Washington. U.S. Pacific NW Forest and Range Expt. Sta. Portland, Ore- gon. 417 pp. Germano, D. J., and C. R. Hungerford. 1981. Reptile population changes with manipulations of Sono- ran Desert shrub. Great Basin Nat. 41:129-138. Heady, H. F., and J. Bartolome. 1977. The Vale re- habilitation program: the desert repaired in southeastern Oregon. USDA For. Serv. Resour. Bull. PNW-70, U.S. Pacific NW Forest and Range Expt. Sta. Portland, Oregon. 139 pp. Kindschy, R. R., and C. Maser. 1978. Jordan Craters research natural area. Federal Research Areas in Oregon and Washington: a guidebook for scien- tists and educators. Suppl. No. 7. Kittleman, L. R. 1973. Guide to the geology of the Owyhee region of Oregon. Museum of Natural History, Eugene, Oregon. Bull. No. 21. 61 pp. Kittleman, L. R., A. R. Green, G. H. Hagood, A. M. Johnson, J. M. McMurray, R. G. Russell, and D. A. Weeden. 1967. Geologic map of the Owyhee region, Malheur County, Oregon. Mu- seum of Natural History, Eugene, Oregon. Bull. No. 8. LoY, W. G., S. Allan, C. P. Patton, and R. D. Plank. 1976. An atlas of Oregon. Univ. of Oregon Press, Eugene, Oregon. 215 pp. Mayhew, W. W. 1968. Biology of desert amphibians and reptiles. Pages 195-356 in S. W. Brown, Jr., ed.. Desert biology. Academic Press, New York. Parker, W. S., and E. R. Pianka. 1976. Ecological ob- servations on the leopard lizard, Crotaphyttis wis- lizeni, in different parts of its range. Herpetolo- gica 32:95-104. Stebbins, R. C. Amphibians and reptiles of western North America. McGraw-Hill, New York. 528 pp. St. John, A. D. 1980. Knowing Oregon reptiles. Salem Audubon Society, 36 pp. Tueller, P. T. 1962. Plant succession on two Artemisia habitat types in southeastern Oregon. Unpub- lished dissertation. Oregon State Univ., Corvallis. Storm, R. M., and R. A. Pimentel. 1949. Herpetological notes from Malheur County, Oregon. Great Basin Nat. 9:59-63. Werschkul, D. F. 1980. The amphibians and reptiles of southeastern Oregon. Unpublished report to the Vale District of the Bureau of Land Manage- ment. 56 pp. PRELIMINARY INDEX OF AUTHORS OF UTAH PLANT NAMES L. Matthew Chatterley,' Blaine T. Welsh,' and Stanley L. Welsh' ,\bstract.— Presented herein is an index to approximately 800 authors of vascular plant names of Utah. A stan- dardized abbreviation is presented for each author. These are listed alphabetically. Following each abbreviation is the full name and birth and death dates (where available) of each individual. In some cases the date of publication is given when biographical information is not known. In compiling a preliminary list of Utah vascular plants (Welsh et al. 1981) an at- tempt was made to standardize abbreviations of author names, but the many inconsis- tencies throughout taxonomic literature sug- gested that additional work was needed. A list of all authors of Utah plant names, com- mon abbreviations, and birth and death dates was begmi. As the material accumulated, the potential usefulness of the information to others working in the field became apparent. There has been no previous single treatment listing the authors of Utah plant names and providing for some consistency of abbrevia- tions. It is hoped that this guide will be a use- ful tool to all those who are concerned with the names of Utah plants. The name of the author of a particular taxon is considered to be a part of the plant name, and is especially useful in citation. The use of the author's name allows one to determine place of pub- lication and other information necessary in taxonomic work. Abbreviations are given first and then the full name. Birth and death dates (where avail- able) are included to aid in correct identi- fication of individuals. In some cases it has been impossible to find biographical informa- tion. We have included only the date of pub- lication for those few individuals. We have chosen to list authors as they are cited in lit- erature. Therefore, two or more authors may be listed together, and one author may ap- pear in several places. This should facilitate easy access to bibliographic citations. We have chosen to apply a double stan- dard of abbreviation in some instances. Ab- breviations of authors' names appearing sepa- rately are allowed longer designations than those where authors names are in com- bination with others (i.e., "T. & G." for Tor- rey and Gray, but "Torr." and "Gray" for the names when they stand alone). Originally, abbreviations were to be based on the most common forms found in liter- ature. Some authors names are sufficiently common in taxonomic literature that their abbreviations have become standard (i.e., Rydb., Wats., and A. Nels.). The names of other authors, often less well known or used only infrequently, were too varied, and it be- came apparent that a general rule was needed, even if it was an arbitrary one. It was not our purpose to rearrange a century of tradition, but strict adherence to a rigid standard was not considered practical. During the process of this compilation the Royal Botanic Gardens, Kew, published the Draft Index of Author Abbreviations: Flower- ing Plants (Halliday et al. 1980). From that list some additional names and dates were ob- tained, but, most importantly, certain criteria for consistent abbreviations were adopted. It is not to be expected that the two lists will correspond exactly. We are only concerned with authors of Utah plant names and, with our center of focus on Utah plants, the scope is therefore much reduced. The general principle followed has been not to abbreviate unless more than two let- ters could be saved by doing so (except where the tradition of taxonomic literature dealing with Utah flora has indicated a common and acceptable alternative; i.e., Marcus Eugene 'Life Science Museum and Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 385 386 Great Basin Naturalist Vol. 42, No. 3 Jones is "Jo^i^s," not "M.E. Jones"). Initials have been added to the abbreviation when more than one author having the same last name occurs on the list. This rule is generally followed, but when a father and son are both listed the son may be designated with an "f" (filius) following the last name. Accents, acutes, umlauts, and other diacritical marks have been omitted. Some names included on the list are not those of taxonomists. Prior to Utah's existence as a state several government exploring expe- ditions made their way through portions of the region. Botanical information was ob- tained during these explorations and pub- lished in government records of the expedi- tions. Authors of those reports are cited in taxonomic literature and are included on this list also (John Charles Fremont, Howard Stansbury, William Emory, John Grubb Parke, Lorenzo Sitgreaves, Stephen Harri- man Long, and others). The work of the late bibliographer of Utah botany. Earl M. Chris- tensen (1967), was especially useful in obtain- ing information on obscure taxonomic pub- lications and authors. Checking of literature has been extensive. The volumes of Taxonomic Literature (Stafleu and Cowan 1976-81) were consulted, as was the three-volume set Biographical Notes upon Botanists (Bamhart 1965). Barnhart's work is a fairly complete reference of taxonomists publishing before the 1940s, and was espe- cially helpful in providing biographical data on older authors. Flora Europaea (Tutin et al. 1964-80), an encyclopedic work on Eu- ropean plants, contains lists of authors and abbreviations at the end of each volume. Those lists were invaluable as sources of names and biographical data on many early European workers. Two lists of abbreviations published in state floras including informa- tion on many major botanical authors were also consulted (Keck 1970, Correll and John- ston 1970). In many cases correct identification of an author was impossible without checking the original citation. Much time was spent con- sulting such standard taxonomic references as Index Kewensis, Union List of Serials, Gray Herbarium Index, and some bibliographic compilations such as Pritzel's Thesaurus Lit- eraturae Botanicae and the four-volume work Index to American Botanical Literature, pub- lished by the Torrey Botanical Club. Some information is still incomplete. Sev- eral older references have not been available to us, some recent author information re- mains sketchy, and, undoubtably, some names have been omitted entirely. Additionally, the discovering of new plant species and the re- naming of others continues to be a vital busi- ness of contemporary taxonomists working with the Utah flora. This guide is, therefore, a preliminary documentation and we solicit help in obtaining corrections and additions. Address your comments to: Herbarium Curator Author Abbreviations M. L. Bean Life Science Museum Brigham Young University Provo, Utah 84602 References Barnhart, J. H. 1965. Biographical notes upon bota- nists. New York Botanical Gardens. 3 Vols. G. K. Hall & Co., Boston. Christensen, E. M. 1967. Bibliography of Utah botany and wildland conservation. BYU Sci. Bull., Biol. Ser. 9(1):1-136. Correll, D. S., and M. C. Johnston. 1970. Abbrevia- tions of authors' names. Pages 1767-1794 in Man- ual of vascular plants of Texas. Contr. Texas Res. Foundation 6:i-1880. Halliday, p., R. D. Meikle, J. Story, and H. Wilkinson. 1980. Draft index of author abbrevia- tions. Royal Botanic Gardens, Kew. Hunt, Rachel M. M. 1958 & 1961. Catalogue of botan- ical books. 2 Vols. Hunt Botanical Library, Pittsburgh. Keck, D. D. 1970. Abbreviations of authors' names. Pages 1551-1576 in P. Munz and D. D. Keck, eds., A California flora. Univ. of California Press, Berkeley. Pritzel, G. a. 1871-1877 (reprint 1972). Thesaurus lit- eraturae botanicae. Otto Koeltz Antiquariat. Stafleu, F. A., and R. S. Cowan. 1976. Taxonomic lit- erature. 4 Vols. Utrecht, The Netherlands. Torrey Botanical Club. 1969. Index to American botanical literature, 1886-1966. 4 Vols. G. K. Hall & Co., Boston. Tutin, T. G., V. H. Heywood, N. A. Burges, D. H. Valentine, S. M. Walters, and D. A. Webb. 1964-80. Flora Europaea. 6 Vols. University Press, Cambridge. 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. September 1982 Chatterley et al.: Authors of Utah Plant Names 387 Index of Authors Abrams Leroy Abrams (1874-1956) Achev Daisy Bird Achey (b. 1906) Aellen Paul Aellen (1896-1973) Agardh Carl Adolf Agardh (1785-1859) Agardh f. Jacob Georg Agardh (1813-1901) Ahles Harry E. Ahles (b. 1924) Airy-Shaw Herbert Kenneth Airy-Shaw (b. 1902) Ait.' William Townsend Aiton (1766-1849) All. Carl AUioni (1725-1804) Al-Shebaz Ihsan A. Al-Shebaz Ames Oakes Ames (1874-1950) L. C. Anderson Loren C. Anderson (b. 1936) Anderss. Nils Johan Andersson (1821-1880) Andrz. Antoni Lukianowicz Andrzejowski (1784-1868) Angstrom Johan Angstrom (1813-1879) Arcangeli Giovanni Arcangeli (1840-1921) Arnold (possibly a pseudonym; ca 1785) Arv.-Touv. Jean Maurice Casimir Arvet-Touvet (1841-1913) Asch. Paul Friedrich Augvist Ascherson (1834-1913) Asch. & Graebn. Paul Friedrich August Ascherson iia34-1913) and Karl Otto Robert Peter Paul Graebner (1871-1933) Asch. & Mag. Paul Friedrich August Ascherson (1834-1913) and Paul Wilhelm Magnus (1844-1914) Asch. & Sweinf. Paul Friedrich August Ascherson (1834-1913) and Sweinf. Ashe William Willard Ashe (1872-1932) Atwood Nephi Duane Atwood (b. 1938) Austin Coe Finch Austin (1831-1880) Babe. Ernest Brown Babcock (1877-1954) Babe. & Stebbins Ernest Brown Babcock (1877-1954) and George Ledyard Stebbins, Jr. (b. 1906) Babington Charles Cardale Babington (1808-1895) Bailey Liberty Hyde Bailey (1858-1954) D. K. Bailey David Kenneth Bailey (b. 1931) V. L. Bailey Virginia Edith Bailey (b. 1908) Baker John Gilbert Baker (1834-1920) Baker Milo Samuel Baker (1868-1961) Baker & Clausen Milo Samuel Baker (1868-1961) and Jens Christian Clausen (1891-1969) Balbis Giovanni Battista Balbis (1765-1831) Ball Carleton Roy Ball (1873-1958) Banks Joseph Banks (1743-1820) Banks & Sprengel Joseph Banks (1743-1820) and Kent Polvcarp Joachim Sprengel (1766-1833) Barbey William Barbey (1842-1914) Barkley Fred Alexander Barkley (b. 1908) T. M. Barkley Theodore Mitchell Barkley (b. 1934) Bameby Rupert Charles Bameby (b. 1911) Barneby & Holmgren Rupert Charles Barneby (b. 1911) and Noel Herman Holmgren (b. 1937) Barratt Joseph Barratt (1796-1882) Barton William Paul Gillon Barton (1786-1856) Batch. Frederick William Batchelder (1838-1911) Batsch August Johann Georg Karl Batsch (1761-1802) Baumg. Johann Christian Gottlob Baumgarten (1765-1843) Baxter Edgar Martin Baxter (b. 1903) Beal William James Beal (18.33-1924) Beaman John H. Beaman (b. 1929) Beauv. Baron Ambroise Marie Francois Joseph Palirot deBeauvois (1752-1820) Bebb M. S. Bebb (1833-1895) Becc. Odardo Beccari (1843-1920) Beckwith Edward Griffin Beckwith (1818-1881) Beetle Alan Ackerman Beetle (b. 1913) Behr Hans Hermann Behr (1818-1904) Bennett Arthur Bennett (1843-1929) L. Benson Lyman David Benson (b. 1909) Benson & Walkington Lyman David Benson (b. 1909) and D. L. Walkington (b. 1930) Benth. George Bentham (1800-1884) Benth. & Hook. George Bentham (1800-1884) and Joseph Dalton Hooker (1817-1911) Bernh. Johann Jacob Bernhardi (1774-1850) Besser Wilbert Swibert Joseph Gottlieb Besser (1784-1842) Bessey Charles Edwin Bessey (1845-1915) Betcke Ernest Friedrich (18i5-1865) Bicknell Eugene Pintard Bicknell (1859-1925) Bieb. Friedrich August Marschall Bieberstein (1768-1826) Bigelow Jacob Bigelow (1787-1879) Blake Sidnev Faye Blake (1892-1959) Blank. Joseph William Blankinship (1862-1938) Bluff & Fingerh. Mathias Joseph Bluff (1805-1837) and Karl Antoine Fingerhuth (1802-1876) Blume Carl Ludwig von Blume (1796-1862) BIytt Mathias Nurnsen Blytt (1789-1862) Boeck. Johann Otto Boeckeler (1803-1899) Boiss. Edmond Pierre Boissier (1810-1885) Boissev. & Davids. Charles Hercules Boissevain (1893-1946) and Carol Davidson Boissevain Charles Hercules Boissevain (1893-1946) J. Boivin Joseph Robert Bernard Boivin (b. 1916) Bolander Henry Nicholas Bolander (1831-1897) Bong. August Heinrich Bongard (1786-1839) F. Boott Francis Boott (1792-1863) W. Boott William Boott (1805-1887) Borbas Vincze von Borbas (1844-1905) Boreau Alexander Boreau (1793-1875) Borkh. Moritz Balthazar Borkhausen (1760-1806) Borner C. J. B. Borner (b. 1880) Bornm. Joseph Friedrich Nicolaus Bornmuller (1862-1948) Botsch. Victor P. Botschantzev (b. 1910) Bowden Wray Merrill Bowden (b. 1914) Brack. William Dunlop Brackenridge (1810-1893) Brand August Brand (1863-1930) Brandegee Townsend Stith Brandegee (1843-1925) M. Brandegee Mary Katharine Brandegee (1844-1920) Branner & Coville John Casper Branner (1850-1922) and Frederick Vernon Coville (1867-1937) A. Br. Alexander Carl Heinrich Braun (1805-1877) Breitung August J. Breitung (b. 1913) Brenckle & Cottam Jacob Frederick Brenckle (b. 1875) and Walter Page Cottam (b. 1894) Brewer William Henry Brewer (1828-1910) Brewer & Wats. William Henry Brewer (1828-1910) and Sereno Watson (1826-1892) Briot Pierre Louis Briot (1804-1888) Briq. John Isaac Briquet (1870-1931) Britt. Nathaniel Lord Britton (1859-1934) Britt. & Br. Nathaniel Lord Britton (1859-1934) and Addison Brown (1830-1913) Britt. & Rose Nathaniel Lord Britton (1859-1934) and Joseph Nelson Rose (1862-1928) 388 Great Basin Naturalist Vol. 42, No. 3 Britt. & Rusby Nathaniel Lord Britton (1859-1934) and Henry Hurd Rusby (1855-1940) Britt. & Shafer Nathaniel Lord Britton (1859-1934) and John Adolph Shafer (1863-1918) B.S.P. Nathaniel Lord Britton (1859-1934), Emerson EUick Stems (1846-1926) and Justus Ferdinand Poggenburg (1840-1893) R. Br. Robert Brown (1773-1858) Buch.-Ham. Francis Buchanan-Hamilton (1762-1829) Buch. Franz Georg PhilUpp Buchenau (1831-1906) Buchholz John Theodore Buchholz (1888-1951) Buckley Samuel Botsford Buckley (1809-1884) Bunge Alexander Andrejewitsch von Bunge (1803-1890) Burgsd. Friedrich August Ludwig Burgsdorf (1747-1802) Burman Johannes Burman (1706-1779) Burman f. Nicolaas Laurens Burman (1734-1793) Butters Frederic King Butters (1878-1945) Butters & Abbe Frederic King Butters (1878-1945) and Ernst Cleveland Abbe (b. 1905) Butters & St. John Frederic King Butters (1878-1945) and Harold St. John (b. 1892) Canby William Marriot Canby (1831-1904) Carr. Elie Abel Carriere (1818-1896) Carruth James Harrison Carruth (1807-1896) Cassidy James Cassidy (1844-1889) Cav. Antonio Jose Cavanilles (1745-1804) Chaix Dominique Chaix (1731-1800) C. & S. Ludolf Karl Adalbert von Chamisso (1781-1838) and Diederich Franz Leonhard von Schlectendal (1794-1866) Cham. Ludolf Karl Adalbert von Chamisso (1781-1838) Chatel. Jean Jacques Chatelain (1736-1822) Chaudhri Mohammad Nazeer Chaudri (b. 1932) Chiov. Emilio Chiovenda (1871-1941) Choisy Jacques Denys Choisy (1799-1859) C. Chr. Car! Frederick Albert Christensen (1872-1942) Chuang & Heckard Tsan-Lang Chuang (b. 1933) and Lawrence R. Heckard (b. 1923) S. Clark Stephen L. Clark (b. 1940) Clausen Jens Christian Clausen (1891-1969) Clokey Ira Waddell Clokey (1878-1950) Clover Elzada Urseba Clover (b. 1897) Clover & Jotter Elzada Urseba Clover (b. 1897) and Mary Lois Jotter (b. 1914) Cockerell Theodore Dru Alison Cockerell (1866-1948) Conrad Solomon White Conrad (1779-1831) Correll Donovan Stewart Correll (b. 1908) Cory Victor Louis Cory (b. 1880) Cosson Ernst Saint-Charles Cosson (1819-1889) Cottam Walter Page Cottam (b. 1894) Coult. John Merle Coulter (1851-1928) Coult. & Nels. John Merle Coulter (1851-1928) and Aven Nelson (1859-1952) Coult. & Rose John Merle Coulter (1851-1928) and Joseph Nelson Rose (1862-1928) Covas Guillermo Covas (b. 1915) Cov. Frederick Vernon Coville (1867-1937) Craig Thomas Theodore Craig (b. 1907) Crantz Heinrich Johann Nepomuk von Crantz (1722-1799) Crepin Francois Crepin (1830-1903) Cronq. Arthur John Cronquist (b. 1919) Cronq. & Keck. Arthur John Cronquist (b. 1919) and David Daniels Keck (b. 1903) Crosswhite Frank Samuel Crosswhite (b. 1940) Curran Mary Katherine Curran (1844-1920) Cyrillo Domenico Maria Leone Cyrillo (Cirillo) (1739-1799) Daniels Francis Potter Daniels (1869-1947) Danser Benedictus Hubertus Danser (1891-1943) Darby John Darby (1804-1877) Darlington Josephine Darlington (b. 1905) Daston J. S. Daston (publ. 1946) Daubs Edwin Horace Daubs (publ. 1965) Daveau Jules Alexandre Daveau (1852-1929) A. Davidson Anstruther Davidson (1860-1932) R. J. Davis Ray Joseph Davis (b. 1895) DC. Augustin Pyramus de Candolle (1778-1841) A. & C. DC. Alphonse Louis Pierre Pyramus de Candolle (1806-1893) and Anne Casimir Pyramus de Candolle (1836-1918) Decne. Joseph Decaisne (1807-1882) Degl. Jean Vincent Yves Degland (1773-1841) Dempster Lauramay Tinsley Dempster (b. 1905) Dempst. & Ehrend. Lauramay Tinsley Dempster (b. 1905) and Friedrich Ehrendorfer (b. 1927) Desf. Rene Louiche Desfontaines (1750-1833) Desmarais Yves Desmarais (publ. 1952) Desr. Louis Auguste Joseph Desrousseaux (1753-1838) Desv. Auguste Nicaise Desvaux (1784-1856) Detl. LeRoy Ellsworth Detling (1909-1967) Dewey Chester Dewey (1784-1867) Dieck Georg Dieck (1847-1925) Diels Friedrich Ludwig Emil Diels (1874-1945) A. Dietr. Albert Gottfried Dietrich (1795-1856) D. Dietr. David Nathaniel Friedrich Dietrich (1800-1888) Dippel Leopold Dippel (1827-1914) D. Don David Don (1799-1841) G. Don Georg Don (1798-1856) Donn James Donn (1758-1813) Dorn Robert D. Dorn (publ. 1977) Dougl. David Douglas (1798-1834) Drejer Solomon Thomas Nicolai Drejer (1813-1842) E. Drew Elmer Reginald Drew (1865-1930) Duchesne Antoine Nicolas Duchesne (1747-1827) Dum-Cours. George Louis Marie Dumont de Courset (1746-1824) Dumort. Barthelemy Charles Joseph Dumortier (1797-1878) Dunal Michel Felix Dunal (1789-1856) Dunn David Baxter Dunn (b. 1917) Dunn & Harmon David Baxter Dunn (b. 1917) and William E. Harmon Durand Elias Magloire Durand (1794-1873) Dur. & Hilg. Elias Magloire Durand (1794-1873) and Theodore Charles Hilgard (1828-1875) Durazzo Ippolito Durazzo (1750-1818) Durieu Michel Charles Durieu de Maisonneuve (1796-1878) Dziek. & Dunn Chester T. Dziekanowski and David Baxter Dunn (b. 1917) Earie W. Hubert Earle (b. 1906) Eastw. Alice Eastwood (1859-1953) D. C. Eaton Daniel Cady Eaton (1834-1895) Eckl. & Zeyh. Christian Frederick Ecklon (1795-1868) and Carl Ludwig Philipp Zeyher (1799-1858) Edwards Sydenham Teast Edwards (1769-1819) Edwin Gabriel Edwin (b. 1926) September 1982 Chatterley et al.: Authors of Utah Plant Names 389 Ehrend. Friedrich Ehrendorfer (b. 1927) Ehrh. Friedrich Ehrhart (1742-1795) Eichler Hansjoerg Eichler (b. 1916) Ell. Stephen Elhott (1771-1830) Ellison William L. Ellison (b. 1923) Elmer Adolph Daniel Edward Elmer (1870-1942) Emory William H. Emory (1811-1887) Endl. Stephan Friedrich Ladislaus Endlicher (1804-1849) Engelm. Georg Engelmann (1809V^884) Engelm. & Bigel. Georg Engelmann (1809-1884) and John Milton Bigelow (1804-1878) Engler Heinrich Gustav Adolph Engler (1844-1930) Engl. & Irm. Heinrich Gustav Adolph Engler (1844-1930) and Edgar Irmscher (1887-1968) Engler & Prantl Heinrich Gustav Adolph Engler (1844-1930) and Karl Anton Eugene Prantl (1849-1893) Erskine David S. Erskine (b. 1900) Ewan Joseph Andorfer Ewan (b. 1909) Farw. Oliver Atkins Farwell (1867-1944) Fassett Norman Carter Fassett (1900-1954) Fedde Friedrich Karl Georg Fedde (1873-1942) Fern. Merritt Lyndon Femald (1873-1950) Ferris Roxana Judkins Ferris (b. 1895) Fisch. Friedrich Ernst Ludwig von Fischer (1782-1854) Fisch. & Mey. Friedrich Ernst Ludwig von Fischer (1782-1854) and Carl Anton Andrievic Meyer (179,5-1855) Fisch. & Trautv. Friedrich Ernst Ludwig von Fischer (1782-1854) and Ernst Rudolph Trautvetter (1809-1889) Fisch., Mey., & Trautv. Friedrich Ernst Ludwig von Fischer (1782-1854), Carl Anton Andrievic Meyer (1795-1855) and Ernst Rudolph Trautvetter (1809-1889) Flous M. Femande Flous (b. 1908) Flowers Seville Flowers (1900-1968) Focke Wilhelm Olbers Focke (1834-1922) Forb. & Hemsl. A. E. E. Forberg (b. 1851) and William Rotting Hemsley (1843-1924) Forbes James Forbes (1773-1861) Forsskal Pehr (Peter) Forsskal (1732-1763) Forster A. Forster (1810-1884) Fosberg Francis Raymond Fosberg (b. 1908) Foum. Eugene Pierre Foumier (1834-1884) Franchet Adrien Rene Franchet (1834-1900) Franco Juao Manuel Antonio do Amaral Franco (b. 1921) Franklin John Franklin (1786-1847) Fraser John Eraser (1750-1811) Frem. John Charles Fremont (1813-1890) Fresen. Johann Baptist Georg Wolfgang Fresenius (1808-1866) Fries Elias Magnus Fries (1794-1878) Fritsch Karl F. Fritsch (1864-1934) Gaertner Joseph Gaertner (1732-1791) Gaertn., Mey. & Scherb. Joseph Gaertner (1732-1791), Bemhard Meyer (1767-1836) and Johannes Scherbius (1769-1813) Galloway Leo A. Galloway (publ. 1975) Gand. Michel Gandoger (1850-1926) Garcke Christian August Friedrich Garcke (1819-1904) Garrett Albert Osbun Garrett (1870-1948) Gates Reginald Ruggles Gates (1882-1962) C. Gay Claude Gay (1800-1873) Gentry Howard Scott Gentry (b. 1903) Geyer Carl Andreas Geyer (1809-1853) Gilib. Jean Emmanuel Gilibert (1741-1814) J. M. Gillett John Montagu Gillett (b. 1918) Glad J. B. Glad (publ. 1971) C. C. Gmel. Carl Christian Gmelin (1762-1837) S. G. Gmel. Samuel Gottlieb Gmelin (1744-1774) Godron Dominique Alexandre Godron (1807-1880) Goldie John Goldie (1793-1886) Goodding Leslie Newton Goodding (b. 1880) Goodman George Jones Goodman (b. 1904) Goodm. & Hitchc. George Jones Goodman (b. 1904) and Charles Leo Hitchcock (b. 1902) Gopp. Heinrich Robert Goppert (1800-1884) Gord. & Glend. George Gordon (1806-1879) and Robert Glendinning (fl. 1844-1858) Gould Frank W. Gould (b. 1913) Gould & Kapadia Frank W. Gould (b. 1913) and Zarir Kapadia (b. 1935) Graebner Karl Otto Robert Peter Paul Graebner (1871-1933) Graham Robert Graham (1786-1845) Grant Alva Day Grant (b. 1920) A. & V. Grant Alva Day Grant (b. 1920) and Verne Edwin Grant (b. 1917) V. Grant Verne Edwin Grant (b. 1917) Gray Asa Gray (1810-1888) Gray, Wats. & Robins. Asa Gray (1810-1888), Sereno Watson (1826-1892) and Benjamin Lincoln Robinson (1864-1935) S. F. Gray Samuel Frederick Gray (1766-1828) Greene Edward Lee Greene (1843-1915) Greenman Jesse Moore Greenman (1867-1951) Greenm. & Roush Jesse Moore Greenman (1867-1951) and Eva Myrtelle Roush (b. 1886) Griffith John William Griffith (1819-1901) Griseb. August Heinrich Rudolf Grisebach (1814-1879) Hackel Eduard Hackel (1850-1926) Haenke Thaddaus Haenke (1761-1817) Halacsy Eugene von Halacsy (1842-1913) Hall Harvey Monroe Hall (1874-1932) H. & C. Harvey Monroe Hall (1874-1932) and Frederic Edward Clements (1874-1945) Hand.-Mazz. Heinrich von Handel-Mazzetti (1882-1940) Hanks & Small Lenda Tracy Hanks (1879-1944) and John Kunkel Small (1869-1938) C. A. Hanson Craig Alfred Hanson (b. 1935) Harrington Harold David Harrington (b. 1903) Hartman Cari Johan Hartman (1790-1849) Harv. & Gray William Henry Harvey (1811-1866) and Asa Gray (1810-1888) Hauman Lucien Hauman (1880-1965) Hausskn. Heinrich Care Haussknecht (1838-1903) Hawksworth & Wiens F. G. Hawksworth (fl. 1964) and Delbert Weins (b. 19.35) Haw. Adrian Hardy Haworth (1768-1833) Hayden Ferdinand Vandeveer Hayden (1829-1887) Heil Kenneth D. Heil (b. 1941) Heimerl Anton Heimerl (1857-1942) Heiser Charles Bixter Heiser (b. 1920) Heller Amos Arthur Heller (1867-1944) Hemsley William Botting Hemsley (184.3-1924) 390 Great Basin Naturalist Vol. 42, No. 3 Henckel Leo Victor Felix Henckel von Donnersmarck (1785-1861) Henderson Louis Fomiquet Henderson (1853-1942) Henrard Jan Theodoor Henrard (b. 1881) L. Henry Louis H. Henry (1853-1903) F. Hermann Frederick Joseph Hermann (b. 1906) Herrmann Johann Herrmann (1738-1800) Herter William Gustav Franz Herter (1884-1958) Hess & Dunn Loyd W. Hess and David Baxter Dunn (b. 1917) Hieron. Georg Hans Emmo Wolfgang Hieronymous (1846-1921) Higgins Larry Charles Higgins (b. 1936) Hildebr. Friedrich Hermann Gustav Hildebrand (1835-1915) Hilend Martha Luella Hilend (b. 1902) Hilend & Howell Martha Luella Hilend (b. 1902) and John Thomas Howell (b. 1903) Hill John Hill (1716-1775) A. S. Hitchc. Albert Spear Hitchcock (1865-1935) Hitchc. & Chase Albert Spear Hitchcock (1865-1935) and Mary Agnes Chase (1869-1963) C. L. Hitchc. Charles Leo Hitchcock (b. 1902) Hitchc. & Maguire Charles Leo Hitchcock (b. 1902) and Bassett Maguire (b. 1904) E. Hitchc. Edward Hitchcock (1793-1864) Hoffm. George Franz Hoffman (1760-1826) H. T. Holm Herman Theodor Holm (1854-1932) A. & N. Holmgren Arthur Hermann Holmgren (b. 1912) and Noel Herman Holmgren (b. 1937) Holmgren, Schultz & Lowrey Arthur Hermann Holmgren (b. 1912), Leila M. Schultz and Timothy K. Lowrey N. Holmgren Noel Herman Holmgren (b. 1937) N. & P. Holmgren Noel Herman Holmgren (b. 1937) and Patricia (nee Kern) Holmgren (b. 1940) P. Holmgren Patricia (nee Kern) Holmgren Holz. John Michael Holzinger (1853-1929) Honck. Gerhard August Honckeny (1724-1805) Hook. & Baker Joseph Dalton Hooker (1817-1911) and John Gilbert Baker (1834-1920) Hook. William Jackson Hooker (1785-1865) H. & A. William Jackson Hooker (1785-1865) and George Amott Walker Amott (1799-1868) Hook. & Grev. William Jackson Hooker (1785-1865) and Robert Kaye Greville (1794-1866) Hopkins Milton Hopkins (b. 1906) Hoppe David Heinrich Hoppe (1760-1846) Homem. Jens Wilken Homemann (1770-1841) Host Nicolaus Thomas Host (1761-1834) House Homer Doliver House (1878-1949) J. T. Howell John Thomas Howell (b. 1903) Howell Thomas Jefferson Howell (1842-1912) Hu & Cheng Shiu-ying Hu (b. 1910) and Ching-yung Joyce Cheng (b. 1919) Hubb. Frederic Tracy Hubbard (1875-1962) Hudson William Hudson (1730-1793) Hulten Oskar Eric Gunnar Hulten (1894-1981) H. B. K. Friedrich Wilhelm Heinrich Alexander vop Humboldt (1769-1859), Aime Jacques Alexandre Bonpland (1773-1858) and Karl Sigismund Kunth (1788-1850) Huth Ernest Huth (1845-1897) Hylander Nils Hylander (1904-1970) litis Hugh Hellmut (b. 1925) Isely Duane Isely (b. 1918) Ives Joseph Christmas Ives (1828-1868) Jacq. Nicolaus Jacquin (1727-1817) James Edwin James (1797-1861) Jarm. A. V. Jarmolenko (1905-1944) Jepson Willis Linn Jepson (1867-1946) Jepson & Bailey Willis Linn Jepson (1867-1946) and Liberty Hyde Bailey (1858-1954) B. L. Johnson B. Lennart Johnson (b. 1909) Johnston Ivan Murray Johnston (1898-1960) J. R. Johnston John Robert Johnston (b. 1880) Jones Marcus Eugene Jones (1852-1934) Juz. Sergei Vasilievic Juzepczak (1893-1959) Kar. & Kir. Grigorij Silic Karelin (1801-1872) and Ivan Petrovik Kirilow (1821 or 1822-1842) Karsten Carl Wilhem Gustav Hermann Karsten (1817-1908) Kaulf. Georg Friedrich Kaulfuss (1786-1830) Kearney Thomas Henry Kearney (1874-1956) Kearney & Peebles Thomas Henry Kearney (1874-1956) and Robert Hibbs Peebles (1900-1956) Keck David Daniels Keck (b. 1903) Kellogg Albert Kellogg (1813-1887) Ker John Bellenden Ker (previously John Gawler) (1764-1842) Kerner R. Anton Joseph Kerner von Marilaun (1831-1898) Kiob. (publ. ca 1770) Kit. Paul Kitaibei (1757-1817) Kittell Marie Teresa Kittell (b. 1892) W. Klein William Klein (publ. 1962) Klotzsch Johann Friedrich Klotzsch (1805-1860) Knerr Ellsworth Brownell Knerr (1861-1942) K. Koch Karl Heinrich Emil Ludwig Koch (1809-1879) Koch Wilhelm Daniel Joseph Koch (1771-1849) Koehne Bernard Adalbert Emil Koehne (1848-1918) Koeler George Ludwig Koeler (1765-1807) Koenig Carl Dietrich Eberhard Koenig (1774-1851) Koen. & Sims Carl Dietrich Eberhard Koenig (1774-1851) and John Sims (1740-1831) Koidz. Genichi Koidzumi (1883-1953) Komarov Vladimir Leontjevic Komarov (1869-1945) Krap. Antonio Krapovickas (publ. 1970) Krause Ernst Hans Ludwig Krause (1859-1942) Kuhn Maximilian Friedrich Adeibert Kuhn (1842-1894) Kukenthal Georg Kukenthal (1864-1955) Kunth Karl Sigismund Kunth (1788-1850) Kuntze Carl Ernst Otto Kuntze (1843-1907) Kunze Gustav Kunze (1793-1851) L'Her. Charles Louis de Brutelle L'Heritier (1746-1800) Lag. Mariano Lagasca Y Segura (1776-1839) Lag. & Rodr. Mariano Lagasca Y Segura (1776-1839) and Jose Demetris Rodriguez (1780-1846) Lam. Jean Baptiste Antoine Pierre de Monnet de Lamark (1744-1829) Lam. & DC. Jean Baptiste Antoine Pierre de Monnet de Lamark (1744-1829) and Augustin Pyramus de Candolle (1778-1841) Lam. & Poir. Jean Baptiste Antoine Pierre de Monnet de Lamark (1744-1829) and Jean Louise Marie Poiret (1755-1834) Lambert Alymer Bourke Lambert (1761-1842) Law,son Charles Lawson (1794-1873) Laxmann Erik G. Laxmann (1737-1796) Ledeb. Carl Friedrich von Ledebour (1785-1851) September 1982 Chatterley et al.: Authors of Utah Plant Names 391 Lehm. Johann Georg Christian Lehmann (1792-1860) Leiberg John Bemhard Leiberg (1853-1913) Lej. Alexandre Louis Simon Lejeune (1779-1858) Lellinger David Bruce Lellinger (b. 1937) Lem. Charles Antoine Lemaire (1801-1871) Leminon John Gill Lemmon (1832-1908) Lepage Ernest Lepage (b. 1905) Less. Christian Friedrich Lessing (1809-1862) H. Levi. Augiistin Abel Hector Leveille (1863-1918) Lewis & Szvv'eykowski Frank Harlan Lewis (b. 1919) and Jerzv Szweykowski (b. 1925) Leysser Friedrick Wilhelni Leysser (1731-1815) Lilj. Samuel Liljeblad (1761-1815) Lindl. John Lindley (1799-1865) Lindl. & Gord. John Lindley (1799-1815) and George Gordon (1806-1879) Lindl. & Paxt. John Lindley (1799-1865) and Joseph Paxton (1803-1865) Lindsay George Edmund Lindsay (b. 1916) Link Johann Heinrich Friedrich Link (1767-1851) L. Carl von Linnaeus (1707-1778) Little Elbert Luther Little Jr. (b. 1907) Litv. Dmitrij Ivanovitsch Litvinov (1854-1929) Lois. Jean Louis August Loiseleur-Deslongchamps (1774-1849) Long Stephen Harriman Long (1784-1864) Loudon John Claudius Loudon (1783-1843) Lour. Joao de Loureiro (1717-1791) Love Askell Love (b. 1916) Love & Love Askell Love (b. 1916) and Doris Love (b. 1918) Lund Peter Wilhelm Lund (1801-1880) Maebr. & Payson James Francis Macbride (1892-1976) and Edwin Blake Payson (1893-1927) Maebr. James Francis Macbride (1892-1976) MacGregor Donald Macgregor (1877-1933) Mack. Kenneth Kent Mackenzie (1877-1934) Mack. & Bush Kenneth Kent Mackenzie (1877-1934) and Benjamin Franklin Bush (1858-1937) Macmillan Conway Macmillan (1867-1929) Macoun John Macoun (1831-1920) Maguire Bassett Maguire (b. 1904) Maguire & Cronq. Bassett Maguire (b. 1904) and Arthur John Cronquist (b. 1919) Maguire & Holmgren Bassett Maguire (b. 1904) and .\rthur Hennann Holmgren (b. 1912) Maguire & Woodson Bassett Maguire (b. 1904) and Robert Everard Woodson (1904-1963) Makino Tomitaro Makino (1862-1957) Malte Malte Oscar Malte (1880-1933) Manetti Guiseppe Manetti (1831-1858) Mansfeld Rudolf Mansfeld (1901-1960) Marcv Randolph Barnes Marcv (1812-1887) Marshall Humphrv Marshall (1722-1801) W. T. Marshall William Taylor Marshall (1886-1957) J. Martin James Stillman (b. 1914) Martin Robert F. Martin (b. 1910) Martins Carl Friedrich Phillipp von Martins (1794-1868) Mason Herbert Louis Mason (b. 1896) Mathias Mildred Esther Mathias (b. 1906) Math. & Const. Mildred Esther Mathias (b. 1906) and Lincoln Constance (b. 1909) Maxim. Carl Johann Maximowicz (1827-1891) Maxon William Ralph Maxon (1877-1948) McClatchie Alfred James McClatchie (b. 1906) McClelland John McClelland (1805-1883) McClintock & Epling Elizabeth May McClintock (b. 1912) and Carl Clawson Epling (1894-1968) McKelvey Susan Delano McKelvey (b. 1883) McVaugh Rogers McVaugh (b. 1909) Medicus Friedrich Casimir Medicus (Medikus) (1736-1808) Merr. Elmer Drew Merrill (1876-1956) Mett. Georg Heinrich Mettenius (1823-1866) Mey. & Scherb. Bemhard Meyer (1767-1836) and Johannes Scherbius (1769-1813) C. A. Mey. Carl Anton Andreevic von Meyer (1795-1855) E. Mey. Ernst Heinrich Friedrich Meyer (1791-1858) F. G. Meyer Frederic Gustav Meyer (b. 1917) Mez Carl Christian Mez (1866-1944) Miehx. Andre Michaux (1746-1802) Miers John M. Miers (1789-1879) Milde Carl August Julius Milde (1824-1871) Miller Phillip Miller (1691-1771) Milliken Jessie Milliken (b. 1887) Millsp. Charles Frederick Millspaugh (1854-1923) Mirbel Charles Francois Brisseau de Mirbel (1776-1854) Moench Conrad Moench (1744-1805) Moore Thomas Moore (1821-1887) Moq. Christian Horace Benedict Alfred Moquin- Tandon (1804-1863) Moretti Giuseppe L. Moretti (1782-1853) Morong Thomas Morong (1827-1894) Morot Louis Rene Marie Francois Morot (1854-1915) Morton Conrad Vernon Morton (1905-1972) Mosher Edna Mosher (Publ. 1915) Muell.-Arg. Jean Mueller-Argoviensis (1828-1896) Muenchh. Otto von Muenchhausen (1716-1774) Muhl. Gotthiff Heinrich Ernest Muhlenberg (1753-1815) Munro William Munro (1818-1889) Munz Philip Alexander Munz (1892-1974) Munz & Klein Philip Alexander Munz (1892-1974) and William McKinley Klein (b. 1933) Murray John Andreas Murray (1740-1791) Nakai Takenoshin Nakai (1882-1952) Nash George Valentine Nash (1864-1921) Necker Noel Martin Joseph de Necker (1729-1793) Nees Christian Gottfried Daniel Nees von Essenbeck (1776-1858) Nees & Mey. Christian Gottfried Daniel Nees von Essenbeck (1776-1858) and Franz Julius Ferdinand Meyen (1804-1840) Neese & Welsh Elizabeth Janet Neese (b. 1934) and Stanley Larson Welsh (b. 1928) A. Nels. Aven Nelson (1859-1952) Nels. & Kennedy Aven Nelson (1859-1952) and Patrick Beveridge Kennedy (1874-1930) Nels. & Maebr. Aven Nelson (1859-1952) and James Francis Macbride (1892-1976) E. Nels. Elias Emanuel Nelson (1876-1949) Nesom G. L. Nesom (publ. 1976) Neuwied Maximilian Alexander Philipp Wied-Neuwied (1782-1867) Nevski Sergei Arsenjevic Nevski (1908-1938) Newberry John Strong Newberry (1822-1892) Nicollet Jean Nicholas Nicollet (1786-1843) Nieuwl. Julius Arthur Nieuwland (1878-1936) 392 Great Basin Naturalist Vol. 42, No. 3 Northstrom & Welsh Terry Edward Northstrom (b. 1945) and Stanley Larson Welsh (b. 1928) J. B. S. Norton John Bitting Smith Norton (1872-1966) Nutt. Tliomas Nuttall (1786-1859) O'Neill Hugh Thomas O'Neill (1894-1969) Oakes William Oakes (1799-1848) Oeder Georg Christian von Oeder (1728-1791) OIney Stephen Thayer Olney (1812-1878) Opiz Philipp Maximilian Opiz (1787-1858) Ortega Casimiro Gomez Ortega (1740-1818) Ortgies Karl Eduard Ortgies (1829-1916) Osterh. George Everett Osterhout (1858-1937) Ottley Alice Maria Ottley (b. 1882) Ownbey Francis Marion Ownbey (b. 1910) G. B. Ownbey Gerald Bruce Ownbey (b. 1916) Pallas Peter Simon von Pallas (1741-1811) Pamp. Renato Pampanini (1875-1949) Parish Samuel Bonsall Parish (1838-1928) Parke John Gnibb Parke (publ. 1855) Parker Kitty Lucille Parker (b. 1910) Pari. Filippo Parlatore (1816-1877) Parodi Lorenzo Raimundo Parodi (1895-1966) Parry Charles Christopher Parry (1823-1890) Pax Ferdinand Albin Pax (1858-1942) Pax & K. Hoffm. Ferdinand Albin Pax (1858-1942) and Karl August Otto Hoffmann (1853-1909) Paxton Joseph Paxton (1801-1865) Payne Willard William Payne (b. 1934) Payson Edwin Blake Payson (1893-1927) Pease & Moore Arthur Stanley Pease (1881-1964) and Albert Hanford Moore (b. 1883) Peck Morton Eaton Peck (1871-1959) Peebles Robert Hibbs Peebles (1900-1956) Pennell Francis Wliittier Pennell (1886-1952) Perry Matthew Calbraith Perry (1794-1858) Pers. Christiaan Hendrick Persoon (1761-1836) Petrak Franz Petrak (1886-1973) Phil. Rudolf Amandus Philippi (1808-1904) Pilger Robert Knud Friedrich Pilger (1876-1953) Piller & Mitterp. Mathias Filler (1733-1788) and Ludwig Mitterpacher von Mitterburg (1734-1814) Pilz G. E. Pilz (publ. 1978) Piper Charles Vancouver Piper (1867-1926) Piper & Beattie Charles Vancouver Piper (1867-1926) and Frederick Steere Beattie (1884-1939) Planchon Jules Emile Planchon (1823-1888) Poir. Jean Louis Marie Poiret (1755-1834) Porsild Alf Erhng Porsild (b. 1901) C. L. Porter Charles Lyman Porter (b. 1889) T. C. Porter Thomas Conrad Porter (1822-1901) Port. & Coult. Thomas Conrad Porter (1822-1901) and John Merie Coulter (1859-1928) Prain David Prain (1857-1944) PrantI Kari Anton Eugene Prantl (1849-1893) Presl Carel Borowag PresI (1794-1852) J. & C. Presl Jan Swatopluk Presl (1791-1849) and Carel Borowag Presl (1794-1852) Pringle James S. Pringle (b. 1937) Pritz. George August Pritzel (1815-1874) Purpus Joseph Anton Purpus (1860-1932) Pursh Frederick Traugott Pursh (1774-1820) Raf. Constantine Samuel Rafinesque-Schmaltz (1783-1840) Ramaley Francis Ramaley (1870-1942) Rattan Volney Rattan (1840-1915) Raup Hugh Miller Raup (b. 1901) Rauschert Stephen Rauschert (b. 1931) Raven Peter Hamilton Raven (b. 1936) Rech. Kari Rechinger (1867-1952) Rech. f. Karl Heinz Rechinger (b. 1906) Red. Pierre Joseph Redoute (1761-1840) Rees Abraham Rees (1743-1825) Regel Eduard August von Regel (1815-1892) Rehder Alfred Rehder (1863-1949) Rehmann Antoni Rehmann (1840-1917) Reichenb. Heinrich Gottlieb Ludwig Reichenbach (1793-1879) Retz. Anders Jahan Retzius (1742-1821) Reveal James Lauritz Reveal (b. 1941) Reveal & Brotherson James Lauritz Reveal (b. 1941) and Jack D. Brotherson (b. 1938) Reveal, Broome, & Beatley James Lauritz Reveal (b. 1941), C. Rose Broome and Janice C. Beatley Rich. Louis Claude Marie Richard (1754-1821) A. Richards Alfred Richardson (publ. 1976) Richards. John Richardson (1787-1865) Richter Karl Richter (1855-1891) Ricker Percy Leroy Ricker (1878-1973) Riddell John Leonard Riddell (1807-1865) J. W. Robbins James Watson Robbins (1801-1879) Robins. Benjamin Lincoln Robinson (1864-1935) Robins. & Fern. Benjamin Lincoln Robinson (1864-1935) and Merritt Lyndon Fernald (1873-1950) Robins. & Greenm. 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Rudd (b. 1910) Ruiz & Pavon Hipolito Ruiz (1754-1815) and Jose Antonio Pavon (1750-1844) Rumpler Theodor Rumpler (1817-1891) Rupr. Franz Josef Ruprecht (1814-1870) Rydb. Per Axel Rydberg (1860-1931) St. John Harold St. John (b. 1892) St.-Yves Alfred Saint-Yves (1855-1933) Salisb. Richard Anthony Salisbury (1761-1829) Sarg. Charies Sprague Sargent (1841-1927) Savi C. Gaetano Savi (1769-1844) J. H. Schaffn. John Henry Schaffner (1866-1939) Schauer Johann Conrad Schaure (1813-1848) September 1982 Chatterley et al.: Authors of Utah Plant Names 393 Scheele George Heinrich Adolf Scheele (1808-1864) Schinz & R. Keller Hans Schinz (1858-1941) and Robert Keller (1854-1939) Schkuhr Christian Schkuhr (1741-1811) Schlecht. Diederich Franz Leonhard von Schlechtendal (1794-1866) Schleicher Johann Christoph Schleicher (1768-1834) Schleiden Matthias Jacob Schleiden (1804-1881) Schneider Camillo Karl Schneider (1876-1951) Schnitzl. Adalbert Carl Friedrich Schnitzlein (1814-1868) Schoener Carol Susan Schoener (b. 1946) Schott Heinrich Wilhelm Schott (1794-1865) Schrader Heinrich Adolph Schrader (1767-1836) Schrank Franz von Paula von Schrank (1747-1835) Schreber Johann Christian Daniel von Schreber (1739-1810) Schultes Josef August Schultes (1773-1831) Schult. & Schult. "josef August Schultes (1773-1831) and Julius Hermann Schultes (1804-1840) Schultz-Bip. Carl Heinrich Schultz-Bipontinus (1805-1867) Schuiz Otto Eugene Schulz (1874-1936) K. Schiim. Karl Moritz Schumann (1851-1904) Schur Philipp Johann Ferdinand Schur (1799-1878) Schweigger August Friedrich Schweigger (1783-1821) Schweigg. & Koerte August Friedrich Schweigger (178.3-1821) and Franz Koerte (1782-1845) Schwein. Ludwig David von Schweinitz (1780-1834) Scop. Giovanni .\ntonio Scopoli (1723-1788) Scribn. Frank Lamson Scribner (1851-1938) Scribn. & Merr. Frank Lamson Scribner (1851-1938) and Elmer Drew Merrill (1876-1956) Scribn. & Sm. 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John Kunkel Small (1869-1938) and Arthur John Cronquist (b. 1919) Smiley Frank Jason Smilev (b. 1880) C. P. Sm. Charles Piper Smith (1877-1955) J. E. Sm. James Edward Smith (17.59-1828) J. G. Sm. Jared Gage Smith (1866-1925) Smyth Bernard Bryan Smyth (1843-1913) Sobol. Gregorius Fedorovitch Sobolevski (1741-1807) Solbrig Otto Thomas Solbrig (b. 1930) Soulange-Bodin Ettienne Soulange-Bodin (1774-1846) Spach Edouard Spach (1801-1879) Spegaz. Carlo Lingi Spegazzini (1858-1926) Sprengel Kurt Polycarp Joachim Sprengel (1766-1833) Stacey John William Stacey (1871-1943) Standley Paul Carpenter Standley (1884-1963) Stansb. Howard Stansbury Stapf Otto Stapf (1857-1933) Staudt Gunther Staudt (publ. 1961) Sternb. Caspar Maria (Graf) von Sternberg (1761-1838) Steudel Ernst Gottlieb von Steudel (1783-1856) Stockwell William Palmer Stockwell (1898-1950) Stokes Susan Gabriella Stokes (1868-1954) Strother John Lance Strother (b. 1941) Sluckey Ronald Lewis Stuckey (b. 1938) Sturm Jakob W. 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Ivar Frederick Tidestrom (1864-1956) Tidestr. & Kittell Ivar Frederick Tidestrom (1864-19,56) and Sister Marie Teresa Kittell (b. 1892) Toft & Welsh Catherine Ann Toft (b. 1950) and Stanley Larson Welsh (b. 1928) Tomb Andrew Spencer Tomb (b. 1943) Torr. John Torrey (1796-1873) Torr. & Frem. John Torrev (1796-1873) and John Charies Fremont (1813-1890) T. & G. John Torrev (1796-1873) and Asa Gray (1810-1888) Torr. & Hook. John Torrey (1796-1873) and William James Hooker (1785-1865) Trautv. Ernst Rudolf von Trautvetter (1809-1889) Trel. William Trelease (1857-1945) Trev. Ludolf Christian Treviranus (1779-1864) Trin. Cari Bernhard von Trinius (1778-1844) Trin. & Rupr. Carl Bernhard von Trinius (1778-1844) and Franz Josef Ruprecht (1814-1870) Turcz. Porphir Kiril Nicolas Stepanovich Turczaninow (1796-1864) Turner Billie Lee Turner (b. 1925) Underw. Lucien Marcus Underwood (1853-1907) Vahl Martin Hendriksen Vahl (1749-1804) Vail Anna Murray Vail (b. 1863) 394 Great Basin Naturalist Vol. 42, No. 3 Van Houtte Louis Van Houtte (1810-1876) Vasey George Vasey (1822-1893) Vasey & Scribn. George Vasey (1822-1893) and Frank Lamson Scribner (1851-1938) Veil. Jose Mariano da Conceicao Velloso (1742-1811) Vent. Etienne Pierre Ventenat (1757-1808) Vill. Dominique Villars (1745-1814) Vilm. Pierre Louis Francois Leveque de Vilmorin (1816-1860) Vitman Fulgenzio Vitman (1728-1806) Voss Andreas Voss (1857-1924) Wagner Warren Herbert Wagner (b. 1920) Wahl. Georg Wahlenberg (1780-1851) Waldst. & Kit. Franz de Paula Adam von Waldstein (1759-1823) and Paul Kitaibel (1757-1817) Wallr. Carl Friedrich Wilhelm Wallroth (1792-1857) Walp. Gerhard Wilhelm Walpers (1816-1853) Walter Thomas Walter (1740-1788) Wangenh. Friedrich Adam Julius von Wangenheim (1747-1800) Ward Lester Frank Ward (1841-1913) Warder John Aston Warder (1812-1833) Waterfall Umaldy Theodore Waterfall (1910-1971) E. E. Watson Elba Emanuel Watson (1871-1936) Wats. Serene Watson (1826-1892) Wats. & Coult. Sereno Watson (1826-1892) and John Merle Coulter (1851-1928) T. J. Watson T. J. Watson (publ. 1977) Webb Jonathan Edwards Webb ?? (publ. 1892) D. A. Webb David Allardice Webb (b. 1912) Weber Georg Heinrich Weber (1752-1828) Weber & Mohr Georg Heinrich Weber (1752-1828) and Daniel Mathias Heinrich Mohr (1779-1808) W. A. Weber William Alfred Weber (b. 1918) Weigel Christian Ehrenfried von Weigel (1748-1831) Weihe & Nees Carl Ernst August Weihe (1779-1834) and Christian Gottfried Daniel Nees von Essenbeck (1776-1858) Weinm. Johann Anton Weinmann (1782-1858) Welsh Stanley Larson Welsh (b. 1928) Welsh & Atwood Stanley Larson Welsh (b. 1928) and Nephi Duane Atwood (b. 1938) Welsh & Bameby Stanley Larson Welsh (b. 1928) and Rupert Charles Bameby (b. 1911) Welsh & Goodrich Stanley Larson Welsh (b. 1928) and Sherel Goodrich (b. 1943) Welsh & Johnston Stanley Larson Welsh (b. 1928) and Barry C. Johnston Welsh & Moore Stanley Larson Welsh (b. 1928) and Glen Moore (b. 1917) Welsh & Reveal Stanley Larson Welsh (b. 1928) and James Lauritz Reveal (b. 1941) Welsh, Atwood, & Reveal Stanley Larson Welsh (b. 1928), Nephi Duane Atwood (b. 1938) and James Lauritz Reveal (b. 1941) Wendl. Hermann A. Wendland (1823-1903) Wheeler George Montague Wheeler (b. 1842) Wherry Edgar Theodore Wherry (b. 1885) Whipple Amiel Wicks Whipple (1818-1863) White Theodore Greely White (1872-1901) Wiegand & Backeberg Karl McKay Wiegand (1873-1942) and Curt Backeberg Wiggers Friedrich Heinrich Wiggers (1746-1811) Wiggins Ira Loren Wiggins (b. 1899) Wight William Franklin Wight (1874-1954) Wikstr. Johan Emanuel Wikstrom (1789-1856) Wilkes Charles Wilkes (1798-1877) Willd. Carl Ludwig von Willdenow (1765-1812) L. O. Williams Louis Otho Williams (b. 1908) Williams Thomas Albert Williams (1865-1900) F. D. Wilson Frank D. Wilson (publ. 1963) Winkler Hubert J. P. Winkler (1875-1941) Wisliz. Friedrich Adolph Wislizenus (1810-1889) With. William Withering (1741-1799) Wittm. Marx Carl Ludewig Wittmack (1839-1929) F. T. Wolf Franz Theodor Wolf (1841-1924) S. J. Wolf & Packer S. J. Wolf and John G. Packer (b. 1929) Wood Alphonso Wood (1810-1881) Woodson Robert Everard Woodson (1904-1963) Wooton Elmer Ottis Wooton (1865-1945) Woot. & Standi. Elmer Ottis Wooton (1865-1945) and Paul Carpenter Standley (1884-1963) Wormsk. Morton Wormskiold (1783-1845) Wulfen Franz Xavier Wulfen (1728-1805) Yates Harris Oliver Yates (b. 1934) Yuncker Truman George Yuncker (1891-1964) Zabel Hermann Zabel (1832-1912) NEST SITE SELECTION IN RAPTOR COMMUNITIES OF THE EASTERN GREAT BASIN DESERT Dwight G. Smith' and Joseph R. Murphy^ Abstract.- Measures of niche breadth and overlap were used to compare nest site selection in a community of 10 raptor species and the Raven nesting in the eastern Great Basin Desert. Three variables were examined: nest site tvpe, elevation, and exposure. Results .suggest a division of component raptor species into relatively abundant core species that .show wide niche breadths and uncommon fringe species with narrow niche breadths. Differences in use of each resource are most pronounced along elevation gradient in which three guilds are evident that correspond to raptor species groupings that nest at higher, middle, and lower elevations. Each guild is comprised of a mix of core and fringe species. Raptor species with highest overlap along one or more nest site variables examined are separated bv differences in activity patterns. In this study we compare nest site selec- tion in a community of 10 raptor species and the Raven {Corvus corax) nesting in the east- em Great Basin desert of Utah. Nest site selection is a function of many variables, including proximity of foraging habitat, protection of nest and young, ther- mal environment of nest, and spatial inter- actions within the community. Nest sites of several raptor species in the eastern Great Basin have some common characteristics and appear similar, suggesting the occurrence of interspecific competition for available nest sites. Observations of occasional appropri- ation of Ferruginous Hawk {Buteo regalis) and Red-tailed Hawk {Buteo iamaicensis) nests by Great Horned Owls {Bubo virgi- nianus); Raven nests by Great Horned Owls; and Great Horned Owl nests by Golden Eagles {Aquih chrysaetos) and Prairie Fal- cons {Falco mexicanus) indicate that nest site availability may be a limiting resource oper- ating during periods of high raptor density (Smith and Mvirphy 1973). Conversely, parti- tioning of nest site resources may reduce competition among raptor species and facil- itate coexistence. We compared raptor ne.st site selection and extent of interspecific overlap using three nest site variables: type of placement, exposure, and elevation. The Raven is in- cluded in the comparison because it is a func- tional raptor, constructs nests that may be appropriated by raptor species, and competes for nesting sites with several raptor species. Study Area Long-term raptor studies began on a 7700 km2 portion of the eastern Great Basin desert in winter 1966-1967. In previous papers we have presented observations on raptor popu- lation dynamics on a smaller 207-km2 in- tensive study area (Smith and Murphy 1973) and described the response of large raptor species to fluctuations of their prey (Smith and Murphy 1979, 1981). Data for this study are from a 1170 km^ segment of our original study area, which includes portions of Utah and Tooele counties in central Utah. Topographically, the area is characterized by broad, flat, alkaline valleys separated by high, north-south oriented hills and ranges. Valley elevations range from 1460 to 1620 m and maximum elevations range from 1830 to 2440 m. Climatically, the area is a northern cold desert (Shelford 1963). Annual precipitation averages 38 cm and monthly temperatures average from -5 C in January to 24 C in July, with wide daily and seasonal variations. Two distinct vegetative associations are present. The desert shrub community occurs over the lower elevations and covers the val- ley floors. It consists of shrubs, herbs, and grasses, several of which form large, homo- 'Department of Biolog\'. Southern Connecticut State College. New Haven, Connecticut 06515. ^Department of ZoologV', Brigham Young University, Provo, Utah 84602. 395 396 Great Basin Naturalist Vol. 42, No. 3 Table 1. Yearly raptor populations nesting in the central Utah study area, 1967-1970. Total Raptor species 1967 1968 1969 1970 nests Golden Eagle 7 13 13 11 44 Ferruginous Hawk 15 28 34 13 90 Red-tailed Hawk 5 10 12 11 38 Swainson's Hawk 2 2 3 1 8 Prairie Falcon 1 1 1 2 5 Kestrel 3 3 2 2 10 Marsh Hawk 0 2 2 3 7 Great Homed Owl 6 14 16 10 46 Short-eared Owl 0 1 1 1 3 Burrowing Owl 1 2 4 3 10 Raven 4 5 5 3 17 geneoas stands under certain edaphic soil conditions. Predominant desert shrub species include big sagebrush (Artemisia tridentata) on the better drained soils and greasewood (Sarcobatus vermiculatus) on the poorly drained valley floors. The well-drained slopes and hills support a dwarf conifer community of Utah juniper (Juniperus osteosperma) and pinyon pine (Pinus monophylla), which occur in stands of widely varying density. A number of abandoned quarries, some of considerable size, are located in the foothills. Many have sheer cliffs ranging from 5-80 m in height. A few structures, all in various states of disrepair, were associated with some quarry sites. These and an abandoned gun- nery tower at the U.S. Army Deseret Depot located in Rush Valley provided artificial nesting sites for some raptor species. Methods The study was conducted from November 1966 through July 1971. To locate nests, we subdivided the study area into 2.56-km2 units, which were systematically searched in a ro- tating sequence at biweekly intervals throughout the nesting season. Foot searches of cliffs, rock outcrops, and juniper stands were supplemented by vehicle searches through desert shrub. Fixed-wing aircraft sur- veys were used two years to find nests but were of limited use due to minimum speed and altitude requirements. Fieldwork each year was from December through August. During this time a minimum of two days per week were spent on the study area, and total fieldwork per year averaged 1640 hours. Ro- tation of search areas throughout the nesting season ensured that all areas were checked several times, and we believe that we suc- cessfully located all raptor nests each year. For each active nest located we recorded (1) elevation above sea level, (2) direction of exposure, and (3) type of placement. Nest site elevation was measured with a portable al- timeter preset at USGS markers located on the study area. Recorded data were grouped in 50 m intervals. The nest site elevation variable permits comparison of how raptor species place their nests above the valley floors over which they hunted. Direction of nest site exposure was determined by orienta- tion of a line projected at 90 degrees from a cliff wall or tree cavity through a nest center and grouped in one of eight equal sub- divisions of the compass. Hillside tree and ground nests were classed by direction of slope. Nests on level ground were not includ- ed in exposure calculations except where nest site entrance orientation was obvious. We recognized 18 nest-site types including 5 cliff, 7 tree, 5 ground, and burrows (Table 2). To compare raptor and Raven nest site se- lection, we used determinations of nest site niche breadth, overlap, and an average com- munity overlap with respect to each of the three nest site variables. Niche breadth of nest site selection was calculated using the standard information theoretic measure presented by Culver (1972): i8i = ^N ij log Njj 2Ni, m, logr where N^ is the abundance of species in cate- gory j and r is the number of categories. The value of /8j may range from 0 to 1. To compare overlap of raptor and Raven nest site selection we used Horn's (1966) overlap index: ih = ^ [f (N, + Nh..) log (N, + Nh,) N„ log 1 '1 jNhj log Nhj] [(N, + Nh) log (Ni + Nh) - N. log N, - Nh log Nh] September 1982 Smith, Murphy: Raptor Nest Site Selection 397 where N,j is the value for species in category j, Nhj the value of species h in category j, N, is the total of values for species in all categories and Nh is the total of values for species h in all categories. Overlap values may range from 0 to 1. Average community overlap of each raptor species within the raptor community was cal- culated using the formula provided by Cody (1974): ^ 2 J n-1 where x represents average community over- lap of a species and all overlap values exclu- sive of aji are smnmed. Dendrograms were constructed using both unweighted (UPGMA) and weighted (WPGMA) pair group cluster analyses (Sokal and Sneath 1963). Resultant dendrograms were similar and we have presented UPGMA in this paper. Results Populations of raptors and ravens nesting on the study area from 1967 to 1970 are pre- sented in Table 1. The Ferruginous Hawk was the most common raptor each year, varying from 13 nesting pairs in 1970 to 34 in 1969. Nesting populations of Great Horn- ed Owls, Golden Eagles, and Red-tailed Hawks were approximately 50 percent small- er: Golden Eagles varied from 7 pairs in 1967 to 13 in 1968 and 1969, Great Horned Owls from 6 in 1967 to 16 in 1969, and Red-tailed Hawks from 5 pairs in 1967 to 12 in 1969. One other large raptor, the Swainson's Hawk {Buteo swainsoni) averaged two nesting pairs per year (range 1-3 pairs). Nesting popu- lations of medium- and small-sized raptors were consistently low throughout the study period. Only one or two pairs of Prairie Fal- cons nested on the study area each year, and the only slightly more abundant Marsh Hawks (Circus cyaneus) nested in three of four study years (1968-70). Short-eared Owls {Asio flammaeus) were the least common raptor and only one nesting pair was foimd from 1968 to 1970. Raven nesting popu- lations varied from 3 pairs in 1970 to 5 in 1968 and 1969. Although 6 of 11 species con- sidered in this study were uncommon, they are characteristic components of nesting rap- tor populations in this portion of the eastern Great Basin. Table 2. Percent distribution of raptor species in 18 nest site type categories.* Nest Site Type GE GHO FH RtH SwH PF MH SpH SeO BuO Ra Cliff sites Quarry 20+ m 5-19 m 5 m Rock outcrop Tree sites Juniper (platform) Juniper (cavity) Pinyon Pine Cliffrose Cottonwood (platform) Cottonwood (cavity) Lombardy Poplar Ground sites Sagebrush Ricegrass Horsebrush Winter Wheat Dry Wash Burrow Sample size (N) 26.3 19.5 — 7.9 22.7 6.5 — 7.9 47.5 43.4 — 36.8 — 2.2 3.2 5.3 4.5 - 24.6 - - 26.0 53.1 21.2 _ 2.2 2.1 7.9 — — 2.1 2.6 — — 1.0 5.3 44 46 7.5 4.3 2.1 90 5.3 38 30.0 10.0 60.0 87.5 12.5 40.0 30.0 20.0 11.5 23.6 5.9 59.0 10.0 - 74.4 12.8 12.8 - 33.3 33.3 33.3 10 20.0 80.0 10 17 ^E = Golden Eagle, GHO = Great Homed Owl, FH = Ferruginous Hawk, RtH = Red-tailed Hawk, SwH = Swainson's Hawk, PF - Prairie Falcon, MH = Marsh Hawk, SpH = Sparrow Hawk (American Kestrel), SeO = Short-eared Owl, BuO = Burrowing Owl, Ra = Raven. 398 Great Basin Naturalist Vol. 42, No. 3 Golden Eagles nested in 4 of 18 (23.5 per- cent) categories, all on cliff sites or rock out- crop (Table 2). Of the large raptors on the study area. Golden Eagles selected the nar- rowest range of nest sites. Comparatively, Ferruginous Hawks and Red-tailed Hawks se- lected the widest variety of nest sites, each using 9 of 18 (47.4 percent) types. Although both species used cliff and tree sites, major differences in type and frequency of use are evident. Over twice as many Red-tailed Hawk nests were located in cliffs (57.9 per- cent vs. 25.6 percent; d = 6.5; P<0.05). Over 55 percent of Red-tail nests were in cliffs greater than 5 m in height, whereas all Ferruginous Hawk nests were located in low cliffs (<5 m) and rock outcrops. Ferruginous Hawks chose tree sites more frequently (58.3 percent compared to 43.3 percent for Red- tailed Hawks), although both species most commonly constructed nests in junipers. Fer- ruginous Hawks but not Red-tailed Hawks also constructed nests on the ground (13.9 percent located in three of the four Great Ba- sin Desert shrub communities available). Great Homed Owls were found in 6 of 18 (13.5 percent) nest-site types, all of which were in cliffs and junipers. Golden Eagles, Prairie Falcons, and Ravens nested exclu- sively in cliff nest site types, and Swainson's Hawks were restricted to juniper and cotton- wood sites. The two smallest raptors on the study area, the American Kestrel {Falco spar- verius) and Burrowing Owl {Athene cunicu- laria) nested only in cavities, the former in cliffs (70 percent) and trees (30 percent) and the latter in burrows. Two other species that used a narrow range of nest site types were the Marsh Hawk and Short-eared Owl. Both selected nest sites in desert shrub communities. Raptor nest site locations in nine elevation categories are presented in Table 3. Although Golden Eagle nest sites were found in eight of nine elevation categories, ranging from 1460 to 1910 m, 78.5 percent were located at elevations greater than 1680 m. On the aver- age, Golden Eagle nests were located at high- er elevations than those of any other raptor in the study area. Great Horned Owls and Red-tailed Hawks were most similar to the Golden Eagle in nest site placement. Great Horned Owl nests were found at all elevations, although most (82.5 percent) were located in elevations greater than 1600 m. Red-tailed Hawk nests were found in 8 of 9 (88.9 percent) elevation categories and also were primarily located at higher elevations. Comparatively, nests of two other large rap- tor species, the Swainson's Hawk and Ferru- ginous Hawk, were restricted to middle and lower elevations. A total of 85.1 percent of all Ferruginous Hawk nests were found be- tween 1511 and 1660 m and none were placed at elevations higher than 1710 m: All Swainson's Hawk nests were found at eleva- tions between 1460-1610 m. Nest sites of Ravens had the highest average elevation. All were above 1711 m and 94.1 percent were at elevations greater than 1761 m. Nest sites of Prairie Falcons were limited to middle eleva- tions from 1611 to 1760 m, but three species, the Marsh Hawk, Short-eared Owl and Bur- rowing Owl nested exclusively in lower ele- Table 3. Percent distribution of raptor species nests in nine elevation categories. Elevation in meters. 1460- 1511- 1561- 1611- 1661- 1711- 1761- 1811- 1861- Number Species* 1510 1560 1610 1660 1710 1760 1810 1860 1910 of nests GE 4.5 0.0 2.3 4.5 22.7 29.5 15.9 6.8 13.6 44 GHO 4.3 4.3 8.7 13.0 21.7 26.1 10.9 4.3 6.5 46 FH 5.3 21.3 45.7 18.1 9.6 0.0 0.0 0.0 0.0 '90 RtH 2.6 5.3 5.3 31.6 23.7 13.2 15.8 2.6 0.0 38 SwH 12.5 25.0 62.5 0.0 0.0 0.0 0.0 0.0 0.0 8 PF 0.0 0.0 0.0 20.0 40.0 40.0 0.0 0.0 0.0 6 MH 57.1 28.6 14.3 0.0 0.0 0.0 0.0 0.0 0.0 7 SpH 0.0 0.0 0.0 10.0 50.0 0.0 30.0 10.0 0.0 10 SeO 33.3 66.6 0.0 0.0 0.0 0.0 0.0 0.0 0.0 3 BuO 30.0 70.0 0.0 0.0 0.0 0.0 0.0 0.0 0.0 10 Ra 0.0 0.0 0.0 0.0 0.0 5.9 29.4 47.1 17.6 17 *GE = Golden Eagle, GHO = Great Homed Owl, FH = Ferruginous Hawk, RtH = Red tailed Hawk, SwH = Swainson's Hawk, PF = Prairie Falcon, MH = Marsh Hawk, SpH = Sparrow Hawk (American Kestrel), SeO = Short-eared Owl. BuO = Burrowing Owl, Ra = Raven. September 1982 Smith, Murphy: Raptor Nest Site Selection 399 vations, below 1610 m for the Marsh Hawk and below 1560 m for the latter two species. A summary of distribution of raptor nests in eight exposure categories is presented in Table 4. Golden Eagles, Great Horned Owls, Ferruginous Hawks and Red-tailed Hawks had exposures in all eight categories, al- though each exhibited higher frequencies for certain exposures. Thus, 57.5 percent of all Golden Eagle nests had exposures from WNW-NNE but 52.5 percent of Great Homed Owl nests had WSW-WNW expo- sures. Exposure preferences of Ferruginous Hawks and Red-tailed Hawks were slightly less definitive, with the former showing high- est ESE nest site exposure and the latter WSW. The two raptor species that nest ex- clusively in cliff and quarry cavities, the Ra- ven and Prairie Falcon, both had maximum WNW nest site exposures. Short-eared Owls exhibited the narrowest range of nest site ex- posure, with all nests at ENE and ESE, but this may reflect the small sample size for this species. Nest site niche breadth.— Niche breadth represents the range or diversity of a species along a resource gradient. Niche breadth values of raptor species along each variable are presented in Table 5. A significant positive correlation between relative abundance of raptor species and niche breadth {(S) was found for each of the nest site variables examined, i.e., type (T), elevation (E), and exposure (X), (r = 0.62, t = 2.36, P <0.05 for /8t; r = 0.63, t = 2.44, P <0.05 for ^Se; and r = 0.72, t = 3.12, P <0.05, for ySx), suggesting that the most com- mon raptor species select the widest range of nesting sites. The Golden Eagle ranked eighth in range of nest site type (fij) selection and tenth in nest site elevation {ft^) and had the widest breadth of nest site exposure selec- tion (^x)- The three other most abundant rap- tors, the Great Homed Owl, Ferruginous Hawk, and Red-tailed Hawk, consistently ranked highest along each variable. If we as- sume that abundance of the large raptor spe- cies reflects their dominance, this is con- sistent with the predictions of Levins (1968) and McNaughton and Wolf (1970) that domi- nant community species occupy broader niches. The American Kestrel ranked sixth or seventh along each variable and showed broader values of nest site type and exposure but not elevation than the average fi values of all species. Another middle-ranked species, the Raven, had a broad )Sx but narrow fi-^ and /?£. It is widely distributed in this part of the Great Basin desert. Five less common raptor species, the Prairie Falcon, Swainson's Hawk, Marsh Hawk, Short-eared Owl, and Burrowing Owl, showed yS values well below the average in each variable. All showed narrow ranges in one or more of the nest site variables examined. Niche overlap.— Niche overlap was cal- culated for all species pairs for each variable. Overlap values provide a measure of the po- tential impact one species may have on an- other. Patterns of overlap and community structure are best illustrated by dendrograms (Cody 1974), which show how species com- binations that closely overlap along a T.\BLE 4. Percent distribution of raptor nests in eight exposure categories. Specie^ WSW WNW NNW NNE ENE ESE SSE SSW Number of nests GE GHO FH RtH SwH PF MH SpH SeO BuO Ra 15.0 17.5 22.5 17.5 7.5 5.0 7.5 7.5 40 29.5 22.7 6.8 2.3 2.3 11.4 6.8 18.2 44 21.6 2.7 8.1 2.7 6.8 27.0 16.2 14.9 74 23.7 10.5 2.6 7.9 5.3 18.4 18.4 13.2 38 12.5 0.0 0.0 25.0 25.0 37.5 0.0 0.0 8 40.0 40.0 20.0 0.0 0.0 0.0 0.0 0.0 5 0.0 42.9 0.0 28.6 28.6 0.0 0.0 0.0 7 10.0 10.0 40.0 0.0 10.0 20.0 10.0 0.0 10 0.0 0.0 0.0 0.0 66.6 33.3 0.0 0.0 3 0.0 0.0 14.3 0.0 0.0 .57.1 14.3 14.3 7 23.5 41.2 11.8 0.0 11.8 5.9 0.0 5.9 17 ^E = Golden Eagle, GHO = Great Homed Owl, FH = Ferruginous Hawk, RtH = Red-tailed Hawk, SwH = Swainson's Hawk. PF = Prairie Falcon, MH = Marsh Hawk, SpH = Sparrow Hawk (American Kestrel), SeO = Short -eared Owl, BuO = Burrowing Owl, Ra = Raven. 400 Great Basin Naturalist Vol. 42, No. 3 resource dimension will form clusters or guilds. Community dendrograms for each nesting site dimension are presented in Fig- ure 1. Each guild represents a group of spe- cies that exploit a specific nest site dimension in a similar manner. Inspection of the nest site type (NST) den- drogram reveals four guilds, the largest (Guild A) containing six species and the smallest just one (Guild D). Subdivisions within the Guild A are, however, readily ap- parent: one shows close similarity of NST se- lection by the Golden Eagle and three other cliff-nesting raptors. Raven, and Prairie Fal- con, and the second subdivision shows a close relationship between Great Horned Owls and Red-tailed Hawks. The frequent appropri- ation of Golden Eagle and Raven nests by Prairie Falcons and Buteo nests by Great Homed Owls contribute a significant per- centage of the overlap and resulting structure of Guild A. Guild B pairs the Ferruginous and Swainson's hawks and illustrates isolating differences between these and the other large raptor species with respect to selection of nest type. The Marsh Hawk and Short-eared Owl both nest in desert shrub and occupy a separate NST guild. The Burrowing Owl is the only raptor species utilizing burrows for nest sites and is the sole member of its guild. Both UPGMA and WPGMA dendrograms show the Ferruginous and Swainson's hawk combination closer to the Marsh Hawk and Short-eared Owl guild than Guild A contain- ing the other large raptors. This is inconsis- tent with observational data: although 14 percent of Ferruginous Hawk nests were ground sites located in desert shrub, another 53 percent were in Juniper and the remain- der on either rock outcrops or artificial struc- tures. All Swainson's Hawk nests were lo- cated in trees. On the basis of overlap values. Ferruginous Hawks actually were closer to the Great Horned Owl-Red-tailed Hawk guild than the Marsh Hawk-Short-eared Owl guild (average pooled 9 values of 0.6994 and 0.3823, respectively). Clustering is maximal in the nest site ele- vation (NSF) dendrogram in which two guilds are evident. Golden Eagles are placed in Guild A, along with Great Horned Owls, Red-tailed Hawks, Prairie Falcons, American Kestrels, and Ravens. Raptor species compo- sition of Guild A is identical to Guild A of NST dendrogram and species composition of Guild B corresponds to NST Guilds B, C, D. The two NSE guilds neatly separate raptor species that nest at middle and low elevations from those that select nest sites at com- paratively higher elevations. NSE Guild B may be further subdivided into a cluster of two raptor species, the Ferruginous and Swainson's hawks, which nested at middle elevations in the knolls and foothills, and the cluster of three raptor species, the Marsh Hawk, Short-eared Owl, and Burrowing Owl, which nested in the valleys. Two guilds are also formed in the nest site exposure (NSX) dendrogram. Raptor species composition of these guilds reveals inter- esting similarities and differences when com- pared to NST and NSE dendrograms pre- viously examined. Differences in NSX preferences split the Ferruginous Hawk- Table 5. Niche breadth parameters of nest site dimensions. Species Number of nests- Nest type Nest elevation /8e Nest exposure Golden Eagle Great Homed Owl FerRiginous Hawk Red-tailed Hawk Swainson's Hawk Prairie Falcon Marsh Hawk American Kestrel Short-eared Owl Burrowing Owl Raven Average (all species) 44 .4581 (8) .8241 (10) .9439(11) 46 .4679 (9) .9043(11) .8625 (8) 90 .5033 (10) .6268 (8) .8870 (9) 38 .6312(11) .8030 (9) .9929 (10) 8 .1280(1) .4097 (3) .6352 (5) 5 .3050 (4) .4801 (5) .5073 (2) 7 .2706 (3) .4351 (4) .5189 (3) 10 .4347 (7) .5317 (6) .7740 (7) 3 .3731 (6) .2897 (2) .3061 (1) 10 .1699(2) .2780 (1) .5551 (4) 17 .3637 (5) .5403 (7) .7422 (6) .3734 .5566 .7023 'Figures in parentheses indicate rank within parameter. 'Sample sizes used in nest site exposure determinations differ as follows: Golden Eagle, 40; Ferruginous Hawk, 74. September 1982 Smith, Murphy: Raptor Nest Site Selection 401 0 0 05 10 Nest Site Type Nest Site Elevation Nest Site Exposure t KI Golden Eagle Raven Prairie Falcon Great Horned Owl Red - tailed Hawk American Kestrel Ferruginous Hawk Swainson ' s Hawk Marsh Hawk Short eared Owl Burrowing Owl Golden Eagle Great Horned Owl Red tailed Hawk Prairie Falcon American Kestrel Raven Ferruginous Hawk Swainson ' s Hawk Marsh Hawk Short eared Owl Burrowing Owl Ferruginous Hawk Red tailed Hawk Great Horned Owl Golden Eagle American Kestrel Raven Prairie Falcon Burrowing Owl Marsh Hawk Short eared Owl Swainson ' s Hawk Fig. 1. Dendrograms resulting from UPGMA cluster analysis of niche overlap matrices of nest site selection di- mensions among raptor species and the Raven. Separate dendrograms are constructed for each nest site lesource dimension. 402 Great Basin Naturalist Vol. 42, No. 3 Swainson's Hawk combination, with the for- mer now included in Guild A containing the other large raptor species and the latter placed in the Marsh Hawk and Short-eared Owl cluster. Two species with very similar NSX requirements are the Raven and Prairie Falcon of Guild A. Both utilize a high pro- portion of cavity nest sites with WNW and NNW exposure. Average community overlap.— Average community overlap (a) within the raptor spe- cies commimity was calculated for each nest site variable (Table 6). Average community overlap provides a method for broadly defin- ing core species that characteristically show high {a>ax) values and fringe species with low average {a 2 z < < . o j 4 i f < ■ ■ < \ I 5 \ A>^^ W ■^ ^ \ i W f — ' J V \ 2- a ■ c •i i ♦ . 2 o 0 < s 2 > < [ 5 5 < n 2 ' — ■ 1 ¥ 5 ? s (D 10 S5 _ .5 o "^ 'x: c § i^ ■5 ^ y ^ -5 December 1982 Allred: Ants of Utah 419 5 o JS 9i lo o N c Camponotus: • Q modoc O @ herculeanus ■ sansabeanus D (3) novaeboracensis < 0 7^ \ • 1 ° ~1\ z D z o 6 - z < C5 B 1 ° ; 2 < \ Z < •^ r >- < N > < 8 ■ < IB o • 'J^i!^ - I • ^ 5 V / ^ o jO " 2 • \ o^ • 1 2 D ^ s i: • < r > 1 • [ z 0 J i. z ID z I < ? 5 9 s s R _ ~ ex s.S '° I c Z Q 3 420 Great Basin Naturalist Vol. 42, No. 4 December 1982 Allred: Ants of Utah 421 ? O s lo a. J. ' ■ ' . o - C Formica: • © fusca 0 gnava ■ @ criniventris O © densiventris < • o > • z 2 i < O • z z < z < < o • • ) ° r^/^-r^ z > >• < 1 * N 0 - • > < O • • < • •8 - © f~^ • • ° ® V o > • • S o ( 3 < r~^ J \ < 3 [ 2 O J 1 — -■ 1 !. z 2» z I < o ( ■ 5 9 s CO R Q u 00 3 ♦ o s s fi ' ' ■ ^ ■ ^ ■ z 2 < m o ■ S 2 ■ 1 z < '^ ]•■ ■ m C (0 •H • JA.'- ^ . O CO 1) i ■^ "^'^-N-, .'-^ K • • jJ (1) OJ io w^ 2 CO c a ■o lU ^^V/*^ >: *N,__^ Cj (D -H CO o; ? ^ •H t>0 4-) c e ^1 ^ CO \ ?) .lu v.^ ti CO CO CJ \ ^ \-/~N £ © o ■ \ z 0 0 I" 1^. • ) o p < s \ ■ = a. r—r < 1 r s7 < ^ \ • < o \ • z < . OJ 5 •■ ;^^ ■ • ■ ..^y^^ml ^ J % — "^ (—^ TsT ^^ -J \ i^ )" V " \ "^ ■ ^ ^^^ o • U J • ■*! • ■ • 2 > ■ m ■ ■ c u z 1 • ■r^2» s ■i! g z « o 8 < 1 > < X < 3; ^ 5 ? s lO « -i 3 ^ ^ij o Is ^ 422 Great Basin Naturalist Vol. 42, No. 4 2 ^ ^ a, o .i a,-- ^■5: >. "^ C 0 0 Cu C 0 ^ o, -5 en 0 0 a; ■^ 0-, 0 ^ OS 51 December 1982 Allred: Ants of Utah 423 -^ ^ Is O oj o .i: Q £ C c U C ^ o 5 ^ o-.S 424 Great Basin Naturalist Vol. 42, No. 4 ♦ o s s P; ' ■ ' z < Z 1 i |\ z < s o 2 • ==0 a Formica: • subnuda O subpolita ■ © subnitens O xerophila 1 3 ) ' < - o >- Of 2 r^ z < \ . o s ■= \ 5 ^ X =' ■ ^4_ • • ^ \' ^ - ^ V ^ 3»^(rjv^ • > s o X i z < a h • • • z < 2^ > o o V \ J z D ■? o o 0 V Od 00 10 R 1 =L O H Oh Q aJ ♦ 2 lO s P; ' ■ Formica: • podzolica o © perpilosa ■ puberula D @ rasilis 0 < O 0 z < 2 B z < Z < z < o ° o 2 2 < 0 A < D - Jo A (^ p ~ - JX ^ • ^ T ■ 1 • • • • r^ 0 V \ 2 J 10 1 ♦ . • z o J J o i ' °° i o < S 1J ' ir- a. 3 J o December 1982 Allred: Ants of Utah 425 ♦ o s 9! ^ i [ • Z 3 1 o 1 z < o ?io. • ® • ® • (0 in 3 3 ) "^ o )j CO m a U p ^ "i "'^"^-vv ...rSVi- \ \ •• W -H g CO 1 ^ ^ to p \ r-J \ -^\ 5 S g Cb?1^ • \ _ • /t57;';v--i-Jj»^ • ^^^^P^ Ht- • 2 • n ■ « 5 r-°. .1, . Jz o: r° 0 V o « 5 > I • * o < ■' s J J ' N — _ ? 0> to n "5 ^ 426 Great Basin Naturalist Vol. 42, No. 4 o \ r^ ® \ • r^ ty Z 0 r \.' •t. i i < 3 < 2 > < [ z I < J ? 5 ? s CO 10 t; _ c ^ ::* OJ w a ^ v: "^ ij -Ci c« -* o n T3 cir C 0; o !^ •— ^ z -4 o a. o -J -o. O 3 December 1982 Allred: Ants of Utah 427 S c g c §S c = ^ .41 o s S to < < z • • z < z < Monoraorium 0 minimum Manica @ mutica Myrmecocystus : ■ flaviceps a mendax ] z 2 &---V .^ >- < K • . o 2 2 \ z I ^ [^« =' • O ] f < o • • < \ ^ ^ /T^, ^. 1 ^ o • ^1 -V-: • V < T N ► i • o ] o • •o z < • • • 2! -1 ~"^ J O c ■ 2. o i 8 ( 1 2) 1 D • 1 2 1 ( o 0 [ 0 ■ c ^0 J • \ z 33 z I < ■ . Sj S J 2 lO R E o § a. O £ CC 2 o j; 1) -« D- C ^ u Tl c o •w -n o c ^2 3 "« J3 = 428 Great Basin Naturalist Vol. 42, No. 4 a. t: ^ S -2 ■K ..^ Q c 3 (M ^^ S D ♦ * S ?i fc ' • IB 1 1 7 8 i\ z < s c □ • Z < 1 Myrmica : • emery ana O incompleta ■ lobicornis a © monticola ■ z i 3 < o N ■ • • • z < h } ■ z I I < I X ~ * I V - 1 °/ SL 6 ° A ^° ^<\ ■ ^ ■ 0 . n 1 > ^ _J-^ \_ u, • a CJ^P^ ■ L ^J .^ cP^^<<;;;i:^ • h a 2 D^ ■ ■ O T t . 1 i < 3 2 < \ 5 z I < . — "-^ ^1 c c o o -a; c a. 3 2 p -S; c i.s ^ a; .22 -a Q S SD December 1982 Allred: Ants of Utah 429 ^ ^ K) to R o ■ C 2 a. 5 . Pogonomyrmex: • occidentalls O owyheei ■ sallnus • • V. ••• — % • oo < • r\r-^ z > < • •• • • • K J • • • • • • • • < < Ci • « • • •• • • •• t z < • • • S.'s J • .« .3 s^ p%^k .••• -v^ -.•• • • • \ 4 > • • • • • •• :• 1 ■ • * ■ • • • • •a. % I g • • r ■ ■■•■ i • • • m • • [ • • • • • •J '^ gt (X* < ' ^ _ ? « S K (O fO rt S15 * 1 w to s lo 1 Pogonomyrmex: • rugosus o © californicus ■ maricopa Pheldole Q @ sitarches < 1 D z ID z i < • • • • z < > V < K < • z < I 0 \ z I o r < C5 • • • < • . 4 \ 4 r^ °5 r y \ y ^\-^ 1 1 % 2- 2 •> i < ID J ■ \ s [ z o J L ^-j: i-o^' . 5 ^ - o "y -5 •- ^ .2 Z i 3 d i 430 Great Basin Naturalist Vol. 42, No. 4 ♦ O S s k f^V J • <\ z < 2 o ^ • • < J •• • . o ' ' S o © • § • ] • H • Jz • • po* », 2 o o 1 < 2 > < • z • I < _- s t 5 ? s ID fe a 3 ^ O ;2 o f1 c a, o CJ o ■s 0) a, XI V5 nS Q o -o ^ c ■2 5 3 D ■5; _^ 5 2 St3 December 1982 Allred: Ants of Utah 431 o o- S -c - s 5 f ? S S t> 6 r^ /— ^ l\ \ 1 z < W^^ y "~^^'. ^ V. "^"'^-N-v-/ ^r^ fv /^-^^ ' T^-# L ^"*5n-/''~*"^ r^-*^ Z ^ y^ ' \ AJ V ^^ 1 J 3<^ \ /\ . J^ § /" ^^ ^ K —J. / r X: 5 \ t ^^ - 'K% r-MM ^ll 1 ^ ~>^ J iB < ^ ^ j^*\ S\j^ r s\ iS^Ss^ ^=7 Jii L N S 1 vs ■> > p: Of 3 2 1 i Ji^ 7 1 Vf z o z < ^ ^l-l 9 ; ? S i:; SI 432 Great Basin Naturalist Vol. 42, No. 4 unpublished data for Utah records, and are not intended as synonyms except in those cases as indicated by Creighton (1950), Smith (1979), and other contemporary workers. These and other names applicable to unpub- lished records and identification labels on specimens in collections examined are given in a separate list at the end of this paper as a means to relate those names to the ones I have used in the body of this report. Where two or more subspecies under a given species have been reported from Utah, they can be separated by characters discussed under the respective species listing in the main body of the paper. In the treatment of species, generic and trivial names are listed alphabetically in the form of a checklist without regard to sub- family designations, which may be ascer- tained by reference to the keys. This facil- itates location of a species where an index is not present, and extensive searching need not be done by the person who is not well versed in its phylogenetic placement. Terminology in the keys to subfamilies, genera, and species has been simplified to the lay person's language as far as possible to avoid frustration for those who are not well versed in taxonomic jargon. A glossary of most of the structures referred to in the keys is included, and many structures are figured. Figures 1 and 9 will help in overall structural orientation (Figure 8 was deleted). Glossary Abdomen: the hindmost of the three major body regions of an insect, situated behind the thorax. Antennal club: last two or three antennal segments that are abruptly enlarged to form clublike ap- pearance, sometimes gradually enlarged to much larger terminal segment. Antennal fossa: pit or base of antenna where it at- taches to head. Clavate: clubbed, gradually thickening toward tip. Clypeus: the broad plate, frequently triangular, situated on the front of the head between the base of the antennae and the mandibles. Depressed: flattened or indented. Epinotum (propodeum of some authors): the posterior major subdivision of the thorax (the second major body part of the ant), actually a structural part of the abdomen anterior to the pedicel, fused to the thorax. Erect hair: a hair projecting at essentially a right angle to the surface of the integument. Eye length: maximum measureable length, usually from ventral to dorsal (anterior to posterior). Facet: one of the small lenses of the compound (large) eye. Femur: the third segment of the leg (first long one from the base). Flexor surface: that surface of the segment of a leg that faces and comes close to another such sur- face when a leg is bent. Fossa: a pit or deep groove. Frontal area: a small triangular plate attached to the dorsal edge of the clypeus, situated between the bases of the antennae. Frontal carina: a distinct ridge or line running up- ward from the clypeus, and separating the cheek and antennal base from the median part of the head. Funiculus: that part of the antenna terminal to the scape (the first greatly elongated segment at- tached to the head) consisting of 8 to 11 segments. Gaster: the globular or ovoid segments of the abdomen posterior to the pedicel. Gena: area of head below and behind posterior margin of eye. Head length: maximum measureable length from ven- tral (anterior) border of clypeus to median dorsal (posterior) border of head, or to line equal with corners if posterior edge of head is concave. Head width: maximum width of head exclusive of com- pound eyes if they extend beyond lateral margins of head. Humeral angle: the shoulder angle of the prothorax. Impressed: shallow depressed area or marking. Integument: the outer covering of the body. Keel: a sharply angled, elevated ridge. Labial palps: the two antennalike appendages from the lower lip situated between the mandibles. Major: a worker of the largest subcaste. Mandible: one of the paired, heavily chitinized and usually toothed processes on the extreme latero- ventral sides of the head. Maxillary palps: the antennalike appendages from the second jaws situated on each side between the mandibles and the labium (lower lip). Mesepinotal suture: the transverse indentation that separates the mesonotum from the epinotum. Mesonotum: dorsal surface of mesothorax. Mesopleura: side of mesothorax. Mesothorax: the second major subdivision of the thorax (the second major part of the ant). Metasternum: the ventral area of the thorax between coxae 3. Minor: a worker of the smallest subcaste. Pedicel: the one or two segments between the thorax and gaster, much reduced in diameter and some- times bearing a scale. Petiole: a single-segmented pedicel, or the first seg- ment of a two-segmented pedicel. Postpetiole: the posterior segment of a two-segmented pedicel. Pronotum: the dorsal surface of the prothorax. Prothorax: the anterior major subdivision of the thorax (the second major part of the ant). Psammophore: two rows of long hairs or bristles, one row on each side of the underside of the head. December 1982 Allred: Ants of Utah 433 SCAPE PRONOTUM MESONOTUM BASE OF PETIOLE SCALE OF PETIOLE FIRST GASTRIC SEGMENT Figs. 1-7. 1, Dorsal view of worker showing major structures (from Wheeler and Wheeler 1963); 2, Sickle-shaped mandibles; 3, Frontal carinae covering antennal insertions, and strongly projected laterally; 4, Frontal carinae not strongly projecting laterally; 5, Triangular mandible with large teeth, the basal tooth offset from other teeth; 6, Basal tooth ahgned with other teeth and with mandibular margin; 7, Basal tooth offset and at angle to mandibular margin. Pubescence: a covering of fine soft hairs, usually lying flat against the integument. Puncta: minute holes or pits in the integument. Punctate: possessing puncta. Ruga: see Wrinkle. Scale of petiole: a scalelike, somewhat oval vertical or angled projection arising from the dorsal surface of the petiole. Scale of scape: thick plate at base of scape. Scape: first greatly elongated segment of the antenna at- tached to the head. Sculptured: pattern of elevations and depressions on the integument. Spinasternal cavity: a minute cavity situated on the ventral side of the metathorax medially between coxae 2 and 3 (can be seen only by removing coxae 2 and 3 on one side). Striae: longitudinal impressed lines. 434 Great Basin Naturalist Vol. 42, No. 4 OCCIPITAL BORDER Figs. 9-20. 9, Frontal view of head of worker showing structures and areas of measurement (from Wheeler and Wheeler 1963); 10, Ventral view of head showing short 4-segmented labial palps and longer 6-segmented maxillary palps; 11, 10-segmented antenna with 2-segmented club; 12, 12-segmented antenna with club of 3 or more segments; 13, Terminal, circular anus fringed with hairs; 14, Subterminal, slitlike anus not fringed with hairs; 15, Scape evenly curved near base; 16, Scape abruptly bent at base; 17, Scape with lobed plate extending one-third along its length; 18, Scape with basal collar or flange; 19, 12-segmented antenna without club; 20, Toothed tibial spur of coxa 3. SuBERECT hair: a small hair that projects from the in- tegument at an angle less than 90 degrees but more than 45 degrees. Tergite: the dorsal plate or surface of a segment, usual- ly applicable to the gaster. Thorax: second major division of ant between head and pedicel, composed of prothorax, mesothorax, and epinotum ("metathorax"). Tibia: the fourth segment of the leg (second long one from the base). Vertex: top of head between and posterior to eyes. Whorls: in concentric rings. Wrinkle (ruga): a small ridge or furrow on the surface. Specimens in the Utah State University collection and of unpublished records of December 1982 Allred: Ants of Utah 435 27 31 28 32 29 33 34 Figs. 21-34. 21, Worker of Camponotus sansabeanus showing lack of constncnon oetween 1st two segments of gaster, well-developed eye, and even convexity of thorax (from Creighton 1950); 22, Worker of Ponera opacior show- ing constriction between 1st two segments of gaster, and poorly developed eye (from Creighton 1950); 23, Worker of Myrmica brevinodis showing attachment of postpetiole to anterior end of gaster, and epinotal spines (from Creighton 1950); 24, Worker of Pogonomymiex occidentalis showing pedicel consisting of two segments, and psammophore on underside of head (from Creighton 1950); 25, Worker of Formica rubicunda showing well-developed scale on pedicel of one segment, and dorsal convexity of thorax (from Creighton 1950); 26, Side of epinotum evenly curved and lack- ing abrupt angle; 27, Epinotum with dorsal, conical elevation; 28, Base of scape with rounded corner and poorly de- veloped flange; 29, Base of scape thin with thin flange; 30, Side of epinotum not evenly curved, but with abrupt angle; 31, Spinastemal cavity; 32, Elevated and fringed lateral lobes of spinastemal cavity; 33, Attachment of post- petiole to dorsal surface of pointed gaster; 34, Base of scape with thick flange. 436 Great Basin Naturalist Vol. 42, No. 4 Acknowledgments 35 39 / 40 36 Figs. 35-42. 35, Postpetiole lacking ventral projec- tion; 36, Postpetiole showing ventral projection; 37. Ventral median notch of clypeus; 38, Ventral margin of clypeus lacking median notch; 39, Clypeus with median flattening next to keel; 40, Clypeus with median area gradually curved to fossa; 41, Notch in apex of clypeus; 42, Anterior projection of ventral median lobe of clypeus. George Knowlton were identified pre- dominantly by George Wheeler, Roy Snell- ing, and Andre Francoeur; those in the Brig- ham Young University collection by Arthur Cole; and those in the University of Utah col- lection by Albert Grundmann and Arthur Cole. Ingham identified his own collections, and verified them by comparison with identi- fied specimens in the University of Utah col- lection. I identified the ants, most of which were in the genus Pogonomymiex, which I collected during my two summers of travel over Utah. I am most grateful for the unselfishness and kindness of George F. Knowlton, who loaned his personal collection records to me, which he had been accumulating for many years as a basis for a third Ants of Utah. He contin- ued to periodically send me identifications of contemporary collections that he sent to Wheeler and Francoeur. Wilford Hansen, curator of the Utah State University collec- tion, loaned all their collection and records. Mary Fors, curator of the University of Utah collection, and Albert Grundmann loaned me all their ant collections and records, and Dr. Grundmann kindly provided me with the re- ports of Charles Ingham on his unpublished studies of ants of southwestern Utah. George Wheeler provided records of Utah collections from his personal collection. Russell Ander- son of Southern Utah State College sent their collection for identification. Andre Fran- coeur corrected the identification of some specimens collected by me and provided in- formation concerning the status of some spe- cies of the large genus Formica. I am grateful to Brigham Young University for funds to collect ants over the state, to study collec- tions at other universities, and for partial sup- port of publication costs of the manuscript. Treatment of Species Under each species are listed its known collection localities in Utah, grouped by county (in boldface type), and its general dis- tribution in the United States. Specific states of occurrence are listed only for those inter- mountain ones immediately adjacent to Utah (Colorado, Arizona, Nevada, Idaho, and Wyoming), which may in certain cases sup- port some records for Utah that are question- able. In the specific locality listings for Utah, the source of the collection record is desig- nated in parentheses for each locality, or for a series of localities listed between two rec- ord sources. Isolated collections may be made of a spe- cies out of its normal range, elevation, or habitat. Such instances may be the result of accidental transport of winged females by di- rectional winds or on vehicles. In most in- stances such individuals or small resulting December 1982 Allred: Ants of Utah 437 colonies are not apt to survive long in unfavorable habitats. Where only one or two records are listed for the state, or its occur- rence in Utah is out of place in its otherwise known range, such should be considered as temporarily questionable. Specimens repre- senting some of these records are not avail- able for examination; hence, the records must remain as tentative until verified by addition- al collections. However, one must realize that any new distribution record must always begin with one collection. Following the Utah records, findings of other workers with reference to specific habi- tat, plant community relationships, and ele- vation are indicated. Finally, collection data taken in my study are given with pertinent field observations. Code for Collection Locality Sources A Dorald M. Allred unpublished records AC Allred and Gole 1979 B Buren 1968 BAD Beck et al. 1967 BY Brigham Young University entomology museum C42 Cole 1942 C56 Cole 1956 C68 Cole 1968 Cr Creighton 1950 F Francoeur 1973 G52 Grundmann 1952 G58 Gnmdmann 1958 Gr63 Gregg 1963 Gr72 Gregg 1972 H Hayward 1945 159 Ingham 1959 163 Ingham 1963 K70 Knowlton 1970 K75 Knowlton 1975 KU George F. Knowlton unpublished records O Olsen 1934 RAU Russell Anderson unpublished records RG Rees and Grundmann 1940 S73 Snelling 1973 S76 Snelling 1976 Sm52 Smith 1952 Sm53 Smith 1953 Sm57 Smith 1957 Sm79 Smith 1979 U University of Utah entomology museum US Utah State University entomology museum W67 Wheeler and Wheeler 1967 W70 Wheeler and Wheeler 1970 Wb Weber 1947 Wi Wilson 1955 Wg Wing 1968 WU George C. Wheeler unpublished records Page Reference for Taxonomic Keys Subfamilies 437 Genera and species of subfamily Ponerinae 438 Genera of subfamily Myrmicinae 438 Species of Aphaenogaster 439 Species of Crematogaster 439 Species of Leptothorax 440 Species of Manica 440 Species of Myrmica 440 Species of Pheidole 441 Species of Pogonomyrmex 442 Species of Solenopsis 443 Species of Stenamma 443 Genera of subfamily Formicinae 444 Species of Acanthomyops 444 Species of Camponotus 445 Species of Formica 446 Species of Lasius 450 Species of Myrmecocystus 451 Genera of subfamily Dolichoderinae 443 Species of Gonomyrma 444 Species of Iridomyrmex 444 Keys to the Identification of Utah Ants Key to Subfamilies of Formicidae Workers (Modified by Cole 1942) 1. Pedicel of two segments (Fig. 24) 2 — Pedicel of one segment (Fig. 25) 3 2(1). Frontal carinae close together, do not cover antennal insertions (one species known from Utah, Neivamyrmex californicus) Ecitoninae — Frontal carinae wide apart, cover antennal insertions (Fig. 3) Myrmicinae 3(1). Caster constricted between first two segments (Fig. 22) Ponerinae — Caster not constricted as above (Fig. 21) 4 4(3). Anus (acidopore) terminal, circular and fringed with hairs (Fig. 13) Formicinae — Anus (acidopore) subterminal and ventral, slit-shaped and not fringed with hairs (Fig. 14) Dolichoderinae 438 Great Basin Naturalist Vol. 42, No. 4 Key to Utah Genera and Species of Ponerinae Workers (Modified from Creighton 1950) 1. Petiole slender, narrower dorsally than ventrally Hypoponera opacior — Petiole robust, as wide dorsally as ventrally 2 2(1). Head has coarse puncta Ponera pennsylvanica — Head has fine puncta Hypoponera opaciceps Key to Utah Genera of Myrmicinae Workers (Modified from Cole 1942, Creighton 1950) 1. Postpetiole attached to dorsal surface of gaster, which is flattened dorsally, convex ventrally, and acutely pointed (Fig. 33) Crernato gaster — Postpetiole attached to medioanterior or ventroanterior end of gaster, which is of usual shape, not as above (Fig. 23) 2 2(1). Antenna 10-segmented, has 2-segmented club (Fig. 11) Solenopsis — Antenna has more than 10 segments; club, when developed, has more than 2 segments (Fig. 12) 3 3(2). Antenna 11-segmented 4 — Antenna 12-segmented 6 4(3). Thorax and petiole lack spines or teeth; pronotum never angular (one species known from Utah, M. minimum) Monomorium — Epinotum has spines or teeth (Fig. 23) 5 5(4). Mesepinotal suture relatively deep (one species known from Utah, F. chamherlini) Formicoxenus — Mesepinotal suture faint or absent Leptothorax 6(3). Laterodorsal part of clypeus elevated as a narrow ridge that forms abrupt sem- icircular boundary at front of antennal fossa (one species known from Utah, T. caespitum) Tetramorium — Laterodorsal part of clypeus not as above 7 7(6). Ants of two distinct sizes, no intermediates between the extremes; antenna has a 3-segmented club that is longer than remainder of funiculus Pheidole — Ants of several sizes, with intermediates between the extremes; antenna lacks distinct club, or, if club present, it is shorter or no longer than remainder of funiculus 8 8(7). Last 3 antennal segments together much shorter than remainder of funiculus, do not form an abrupt club, although they may gradually enlarge to the terminal largest segment 9 — Last 3 antennal segments together form an abrupt club nearly as long as remainder of funiculus Leptothorax 9(8). Dorsum of thorax has a relatively deep mesepinotal suture; psammophore ab- sent; head, thorax and petiole sometimes with widely spaced, deep parallel wrinkles 10 — Dorsum of thorax lacks such a suture or at most has a slight impression; psammophore present (Fig. 24); head, thorax and petiole not as above Pogonomyrmex December 1982 Allred: Ants of Utah 439 10(9). Posterior tibial spurs comblike (Fig. 20) 11 — Posterior tibial spurs lack teeth 12 11(10). Epinotum has spines (Fig. 23) Myrmica — Epinotum lacks spines Manica 12(10). Eyes small, poorly developed (Fig. 22); clypeus has 2 keels Stenamma — Eyes large, well developed (Fig. 21); clypeus lacks keels 13 13(12). Spines of epinotum long and narrow, about as long as petiole; head not longer than broad (one species known from Utah, V. lobognathus) Veromessor — Spines of epinotum short and robust, shorter than petiole; head longer than broad Aphaenogaster Key to Utah Species of Aphaenogaster Workers (Modified from Cole 1942, Creighton 1950) 1. Scape of larger workers (sometimes smaller ones) surpasses corner of head by amount less than length of 1st two funicular segments 2 — Scape of large and small workers surpasses corner of head by amount greater than length of 1st two funicular segments 3 2(1). Epinotal spines prominent; body chestnut brown occidentalis — Epinotal spines small tubercles; head and thorax yellowish red, gaster dark brown or black uinta 3(1). Epinotum lacks teeth or spines boulderensis — Epinotum has teeth or spines huachucana Key to Utah Species of Crematogaster Workers (Modified from Creighton 1950, Buren 1968) 1. Postpetiole lacks median groove minutissima — Postpetiole divided by distinct median groove 2 2(1). Dorsum of thorax lacks erect hairs, or has not more than 8 confined to humeral angle of pronotum; rarely has 3 or 4 short hairs in mesepinotal suture 3 — Dorsum of thorax has at least 15 scattered erect hairs 7 3(2). Thorax densely punctate; has one or no erect hairs on humeral angle of pronotum 4 — Thorax not punctate, or, if so, has 2 or more erect hairs on humeral angle of pronotum 5 4(3). Thorax lacks erect hairs depilis — Thorax has one erect hair on humeral angle of pronotum noctuma 5(3). Lower mesopleura has distinct striae 6 — Lower mesopleura lacks striae hespera 6(5). Head completely covered with striae and puncta coarctata — Head smooth and shining at least behind eye, sometimes has weak impressed lines elsewhere mormonum 7(2). Hairs on thorax short, 4 or more on each pronotal shoulder, 2 or more at rear of mesonotum emeryana — Hairs on thorax long, 1 to 3 on each pronotal shoulder; mesonotum lacks hairs cerasi 440 Great Basin Naturalist Vol. 42, No. 4 Key to Utah Species of Leptothorax Workers (Modified from Creighton 1950) 1. Antenna ll-segmented 2 — Antenna 12-segmented 3 2(1). Scape has abundant erect hairs hirticornis — Scape lacks erect hairs, or has only a few 9 3(1). Anterior part of 1st gastric tergite covered with striae and puncta silvestrii — Anterior part of 1st gastric tergite smooth and shining 4 4(3). Posterior half of head mostly smooth and shining; broad central strip that extends forward to antennal lobes lacks sculpture nitens — Head largely or entirely sculptured, its entire surface opaque or feebly shining 5 5(4). Dorsum of thorax densely and evenly punctate, wrinkles absent or feeble; sides of thorax have heavy puncta that obscure wrinkles 6 — Dorsum of thorax has puncta interrupted by prominent wrinkles on at least epinotum and mesonotum; wrinkles on sides of thorax not obscured by puncta 7 6(5). Epinotal spines reduced to short, stumpy angles andrei — Epinotal spines well developed nevadensis 7(5). Scale of petiole has feeble wrinkles and dense puncta furunculus — Scale of petiole has coarse wrinkles 8 8(7). Dorsum of thorax completely covered with coarse longitudinal wrinkles except for small heavy puncta on mesonotum nevadensis — Wrinkles on dorsum of thorax largely confined to epinotum and rear of mesonotum; anterior prothorax has puncta only tricarinatus 9(2). Clypeus lacks ridges, its center usually flattened; mesepinotal suture present and deeply depressed 10 — Clypeus has one or more small median ridges; mesepinotal suture absent, or, if present, is not deep 11 10(9). Erect body hairs long, numerous, usually pointed crassipilis — Erect body hairs short, sparse, usually thickened at tip muscorum 11(9). Longitudinal wrinkles of head delicate, not much coarser than sculpture between them ambiguus — Longitudinal wrinkles of head coarse, notably heavier than sculpture between them rugatulus Key to Utah Species of Manica Workers (Modified from Creighton 1950) 1. Postpetiole has conspicuous ventral projection that extends anteriorly (Fig. 36) hunteri — Postpetiole lacks such a projection (Fig. 35) mutica Key to Utah Species of Myrmica Workers (Modified from Cole 1942, Creighton 1950) 1. Scape gradually and evenly bent at base, upper area does not form right angle at bend (Fig. 15) 2 — Scape abruptly bent at base, upper area forms right angle (Fig. 16) 3 December 1982 Allred: Ants of Utah 441 2(1). Lateral margin of frontal carina strongly angular above antennal insertion, thick and deflected downward toward base of antenna (Fig. 3) incompleta — Lateral margin of frontal carina rounded and thin, deflected upward (Fig. 4) .... brevispinosa 3(1). Bend of scape has large, thick lobed plate that extends backward along basal third of scape (Fig. 17) monticola — Bend of scape has small transverse plate, or thin scale, which surrounds bend like a collar and does not extend backward (Fig. 18) 4 4(3). Ventral surface of postpetiole flat or nearly so, does not form projection in front (Fig. 35) americana — Ventral surface of postpetiole convex, or forms prominent anterior projection (Fig. 36) 5 5(4). Scale of scape forms high, semicircular welt that surrounds scape at bend .. hamulata — Scale of scape not as above 6 6(5). Scale of scape small and diagonally transverse on upper surface of scape, con- tinues as prominent transparent flange along inner surface of scape below bend emeryana — Scale of scape not as above 7 7(6). Epinotal spines bent downward; thorax reddish yellow, head and gaster black with reddish tinge emeryana — Epinotal spines straight, color combination not as above lobicornis Key to Utah Species of Pheidole Workers (Modified from Creighton 1950) 1. Scape of major reaches or surpasses corner of head 2 — Scape does not reach corner of head 3 2(1). Eye has more than 20 facets; head unicolored, dark brown to black desertorum — Eye has less than 15 facets; dorsal surface of head bicolored grundmanni 3(1). Scape of major bent laterally at base toward midline of head, its flattened basal portion as wide as distal part hyatti — Scape not bent at base, or, if slightly so, flat part at base not as wide as distal part 4 4(3). Top of head above eye and usually front face of major marked with elevations or depressions, surface opaque or feebly shining 5 — Top of head not so marked except for hair pits; surface strongly shining 7 5(4). Humeral angles of pronotum of major feebly developed, do not form lateral knobs sitarches — Humeral angles strongly developed into epauletlike lateral knobs 6 6(5). Postpetiole of major lens-shaped, lateral projections well developed pilifera — Postpetiole four-sided, lateral projections absent or poorly developed californica 7(4). Head of major at least 2 mm long virago — Head of major not over 1.5 mm long 8 442 Great Basin Naturalist Vol. 42, No. 4 8(7). Sculpture on head of major extends to top of head, only the corners above the eyes smooth and shining ceres — Sculpture largely confined to anterior half of head, all the posterior half smooth and shining 9 9(8). Mesonotum of major depressed below adjacent portion of pronotum to form distinct step or angular projection between pronotum and mesonotum dentata — Mesonotum not so depressed, forms evenly curved outline with pronotum bicarinata Key to Utah Species of Pogonomyrmex Workers (Modified from Cole 1968) 1. Mandible has 6 teeth, the basal one much reduced; eye situated below approx- imate center of side of head; epinotal spines short, blunt, laterally compressed, joined posteriorly by transverse keel imberbiculus — Mandible has 7 teeth, basal one not reduced; eye at about center of side of head; epinotal spines not as above 2 2(1). Viewed dorsally, lateral lobe of clypeus in front of base of antenna forms ante- riorly-projecting broad blunt process; head much broader than long; eye small, in front view does not extend beyond side of head; 1st segment of gaster broader than long rugosus — Viewed dorsally, lateral lobe of clypeus does not extend anteriorly; head not broader than long or only slightly so; eye large, extends beyond side of head; 1st segment of gaster not broader than long 3 3(2). Base of scape strongly enlarged, broad, its basal flange thick (Fig. 34); wrinkles on head above eye not in concentric whorls; dorsum of postpetiole not longer than broad; epinotal spines always present 4 — Base of scape weakly enlarged, its basal flange thin (Fig. 29); wrinkles of head above eye in concentric whorls; dorsum of postpetiole longer than broad; epinotal spines absent or present 7 4(3). Spaces between wrinkles on head opaque, densely and strongly punctate, producing beaded appearance 5 — Spaces between wrinkles subopaque or shining, not densely or strongly punc- tate, do not produce beaded appearance; spines may be reduced to tubercles ... subnitidus 5(4). Basal tooth of mandible offset, meets short basal mandibular margin at pronounced angle (Fig. 7), or forms a gradual curve with margin occidentalis — Basal tooth not offset, meets long basal mandibular margin at straight angle (Fig. 6) 6 6(5). Dorsum of petiole and postpetiole has numerous strong, wavy, closely spaced, subparallel, usually transverse wrinkles; upper edge of base of scape broadly rounded, thin collar does not project beyond rounded corner (Fig. 28) salinus — Dorsum of petiole and postpetiole not as above, but may have irregular wrinkles or lines; base of scape not broadly rounded, may be angled, thin collar projects beyond upper corner owyheei 7(3). Puncta of head and side of epinotum deep, spaces between wrinkles subopaque maricopa — Puncta of head and thorax weak or absent; spaces between wrinkles strongly shining californicus December 1982 Allred: Ants of Utah 443 Key to Utah Species of Solenopsis Workers (Modified from Creighton 1950) 1. 2nd segment of funiculus at least IV2 times as long as broad 2 — 2nd segment of funiculus at most only slightly longer than broad 3 2(1). Distance between eye and base of mandible IV2 times maximum diameter of eye xyloni — Distance between eye and base of mandible at least 2 times maximum diameter of eye aurea 3(1). Puncta of head much larger in diameter than hairs that arise from them salina — Puncta of head barely larger than hairs that arise from them molesta Key to Utah Species of Stenamma Workers (Modified from Snelling 1973) 1. Median lobe of clypeus extends beyond clypeal ventral margin, in frontal view its apex rounded or truncate (Fig. 42); eyes large, distance between eye and mandible less than 2 times eye length 2 — Median lobe of clypeus does not extend beyond clypeal anterior margin, its apex notched (Fig. 41); eye small, distance between eye and mandible more than 2 times eye length 3 2(1). Eye large, has 8-12 facets in greatest diameter smithi — Eye small, has 6-7 facets in greatest diameter chiricahua 3(1). Frontal carinae of median lobe of clypeus subparallel or slightly divergent apically, area between carinae concave, which gives apex notched appearance 4 — Frontal carinae of clypeus strongly divergent from base, area betwen them flattened, apex not notched huachucanum 4(3). Sides of mesothorax have dense puncta; pronotal sides have coarse puncta between wrinkles occidentale — Sides of mesothorax have coarse wrinkles; pronotal sides lack pimcta between wrinkles diecki Key to Utah Genera of Dolichoderinae Workers (Modified from Cole 1942, Creighton 1950) 1. Scale of petiole well developed (Fig. 25) 2 — Scale of petiole vestigial or absent (one species known from Utah, T. sessile) Tapinoma 2(1). Epinotum has conical elevation (Fig. 27) Conomyrma — Epinotum lacks conical elevation 3 3(2). Dorsum of thorax lacks impression at mesepinotal suture; workers of several sizes (one species known from Utah, L. occidentale) Liometopum — Dorsum of thorax has slight to moderate impression at mesepinotal suture; workers of one size 4 4(3). Erect body hairs long and sparse, absent on scape and tibia Iridomyrmex — Erect body hairs short and numerous, present on scape and tibia (one species known from Utah, F. foetidus) Forelius 444 Great Basin Naturalist Vol. 42, No. 4 Key to Utah Species of Conomyrma Workers (Modified from Creighton 1950) 1. Head and thorax deep reddish yellow, gaster brownish black bicolor — Color not as above insana Key to Utah Species of Iridomyrmex Workers (Modified from Creighton 1950) 1. Scape extends beyond corner of head by amount equal to or greater than length of 1st funicular segment humilis — Scape extends beyond corner of head by amount less than length of 1st funicular segment pruinosus Key to Utah Genera of Formicinae Workers (Modified from Creighton 1950) 1. Antenna has 9 segments (one species known from Utah, B. depilis) Brachymyrmex — Antenna has 12 segments (Fig. 19) 2 2(1). Dorsum of thorax in profile evenly convex (Fig. 21) Camponotus — Dorsum of thorax distinctly depressed behind level of mesonotum (Fig. 25) 3 3(2). Mandible sickle-shaped, inner border has tiny teeth (Fig. 2); maxillary palp 4-segmented, labial palp 2-segmented (one species known from Utah, P. breviceps) Polygerus — Mandible triangular, inner border has large teeth (Fig. 5); maxillary and labial palps not as above 4 4(3). Maxillary palp short, 3-segmented Acanthomyops — Maxillary palp longer, has at least 5 apparent segments (Fig. 10) 5 5(4). Psammophore present (Fig. 24); maxillary palp longer than head Myrmecocystus — Psammophore absent; maxillary palp not longer than head 6 6(5). Frontal carinae prominent, their lateral margins slightly deflected upward; ocelli large and conspicuous Formica — Frontal carinae poorly marked, their lateral margins flat; ocelli small and indistinct or absent 7 7(6). Scape extends past corner of head by at least one-third length of scape, usually more; erect body hairs coarse, long, usually brown or black (one species known from Utah, P. parvula) Paratrechina — Scape does not surpass comer of head; erect body hairs fine, short, and golden . Lasius Key to Utah Species of Acanthomyops Workers (Modified from Wing 1968) 1. Erect hairs on dorsum of gaster segments restricted to or concentrated near posterior margins; in side profile top of scale sharp to moderately sharp; top of scale usually with median indentation interjectus — Erect hairs on dorsum of gaster segments uniformly distributed over entire surface; scale variable in shape 2 December 1982 Allred: Ants of Utah 445 2(1). In side profile, top of scale moderately to greatly blunt; throat has erect hairs from front to rear 3 — In side profile, top of scale moderately to greatly sharp; throat has erect hairs only on one-half to three-fourths of posterior surface 4 3(2). Erect hairs more numerous on dorsum of epinotum than on pro- and mesothorax; 1st femur has 10 or less erect hairs; scape lacks erect hairs murphyi — Erect hairs about evenly distributed over entire dorsum of thorax; 1st femur has 12 or more erect hairs; scape may have or lack erect hairs latipes 4(2). Side of 2nd segment of gaster densely covered with hairs, the distance between hairs less than one-third their length; appendages and most of body densely covered with hairs occidentalis — Side of 2nd segment of gaster moderately to thinly covered with hairs, the dis- tance between hairs more than one-half their length; appendages and body moderately to thinly covered with hairs 5 5(4). Body color yellow; erect hairs on throat 0.13 mm or more in length, those on gaster 0.18 mm or more; hairs delicate creightoni — Body color yellowish brown to brown; erect hairs on throat 0.12 mm or less in length, those on gaster 0.17 mm or less; hairs coarse coloradensis Key to Utah Species of Camponotus Major Workers (Modified from Creighton 1950, Gregg 1963) 1. Ventral border of clypeus depressed in middle to form thin anterior edge, usu- ally with narrow median notch (Fig. 37); clypeus usually has short triangular depression behind notch 2 — Ventral border of clypeus not depressed, edge wide, sometimes with notch; short triangular depression absent 5 2(1). Mesepinotal suture broadly impressed, involves rear of mesonotum and front of epinotum hyatti — Mesepinotal suture not impressed, or, if slightly so, only as groove on front of epinotum 3 3(2). Corner of head strongly shining, sides of head notably narrowed at level of mandibles essigi — Corner of head opaque or feebly shining, sides of head not unusually narrowed at level of mandibles 4 4(3). Clypeus much broader than long nearcticus — Clypeus only slightly broader than long rasilis 5(1). Clypeus lacks distinct ridge or keel, but sometimes has slight one; scape not flattened at base; head broader than long 6 — Clypeus has distinct ridge or keel, sometimes reduced; if reduced, scape flattened at base; head as long as or longer than broad 9 6(5). Scape has scattered erect hairs; entire ant jet black and shining, often with bluish reflections laevigatus — Scape lacks erect hairs except for small cluster at extreme tip; color not as above, but if black then not shining and lacks bluish luster 7 7(6). Scape reaches only to or barely extends beyond corner of head herculeanus — Scape extends beyond corner of head by amount greater than maximum diameter of scape 8 446 Great Basin Naturalist Vol. 42, No. 4 8(7). Pubescence on gaster absent or fine and sparse; entire surface of gaster shining novaeboracensis — Pubescence on gaster coarse and relatively dense, surface dull except for narrow lighter-colored band at posterior edge of each segment modoc 9(5). Scape extends beyond corner of head by amount equal to or greater than length of 1st funicular joint 10 — Scape does not extend beyond corner of head, or, if so, by amount less than length of 1st funicular joint sansaheanus 10(9). Scape distinctly flattened at base, flattened portion forms small lateral lobe semitestaceus — Scape not flattened at base, or, if flattened, lacks lateral lobe 11 11(10). Cheek strongly shining, has tiny inconspicuous puncta ocreatus — Cheek dull or feebly shining, has coarse and conspicuous puncta vicinus Key to Utah Species of Formica Workers (Modified from Creighton 1950, Francoeur 1973, Wheeler and Wheeler 1977) 1. Ventral margin of clypeus has median notch (Fig. 37); short hairs on gaster dense; body bicolored with head and thorax reddish brown or reddish yellow, gaster brown or black; in side profile dorsomedian area of the epinotum between mesoepinotal suture and petiole angled, not broadly curved 2 — Lacks above combination of characters (Fig. 38) 10 2(1). Dorsum of 1st segment of gaster strongly shining, its thin hairs do not obscure delicate rough sculpture 3 — Dorsum of 1st segment of gaster opaque or feebly shining, its dense hairs partially obscure fine leatherlike sculpture 4 3(2). Basal face of petiole has many long erect hairs; other body hairs long and numerous perpilosa — Basal face of petiole lacks long erect hairs, or has cluster of short erect ones near junction with sloping face; other body hairs short and not abundant manni 4(2). Dorsum of thorax lacks erect hairs, or has few fine short inconspicuous ones only on pronotum 5 — Dorsum of pronotum and mesonotum has conspicuous erect hairs 8 5(4). In front face view of largest workers, outer margin of eye reaches or surpasses margin of head 6 — In front face view of largest workers, distinct space evident between outer margin of eye and margin of head 7 6(5). Scape slender, not thickened at tip; basal face of epinotum lacks transverse impression pergandei — Scape robust, tip thickened; basal face of epinotum has distinct transverse impression emeryi 7(5). Scale of petiole has blunt crest; gaster plain brown; upper face of epinotum at right angle to base subintegra — Scale of petiole has sharp crest; gaster blackish brown; upper face of epinotum at greater than right angle to base subnuda December 1982 Allred: Ants of Utah 447 8(4). Gaster has long, stout, silvery, erect hairs blunt at tip; erect hairs on other parts of body about as abundant as those on gaster obtusopilosa — Erect hairs on gaster yellow, not blunt at tip; erect hairs on other parts of body much more sparse than on gaster 9 9(8). Scape has abundant suberect hairs, those on inner surface near tip distinctly erect puberula — Scape lacks suberect or erect hairs wheeleri 10(1). From posterior profile, upper surface of side of epinotum evenly curved to base (Fig. 26); body surface shining 11 — From posterior profile, upper surface of side of epinotum angled before its base (Fig. 30); body surface dull 14 11(10). Scale more than VA length of head; frontal carinae diverge dorsally; posterior face of petiole scale convex pallidefulva — Scape less than VA length of head; frontal carinae subparallel, do not diverge dorsally; posterior face of petiole scale not convex 12 12(11). Scape has short, delicate, whitish, erect hairs lasioides — Scape lacks erect hairs except for small cluster at extreme tip 13 13(12). Thorax has numerous erect hairs neogagates — Thorax lacks erect hairs, or has only 1 or 2 limata 14(10). Dorsal border of head of larger workers strongly concave; pronotum in profile angled between base and upper surface opaciventris — Dorsal border of head of larger workers straight or only slightly concave; pronotum in profile evenly convex 15 15(14). Erect hairs on pronotum distinctly broader at tip than at base 16 — Erect hairs on pronotum absent, or, if present, taper from base to pointed tip or of equal width throughout their length 20 16(15). Tibia has erect or suberect hairs in addition to double row on flexor surface; scape hairs variable, often erect microgyria — Tibia lacks erect hairs except for double row on flexor surface; scape lacks erect hairs except for few at extreme tip 17 17(16). Dorsal border of head evenly convex querquetulana — Dorsal border of head flat or concave 18 18(17). Crest of petiole lacks erect hairs; pubescence on dorsum of gaster thin, does not wholly conceal surface at rear edges of segments whymperi — Crest of petiole has erect hairs; pubescence on dorsum of gaster dense, wholly conceals surface 19 19(18). Erect hairs on dorsum of head and thorax sparse, or inconspicuous or absent .... rasilis — Erect hairs on dorsum of head and thorax abundant and conspicuous densiventris 20(15). Body bicolored with head and thorax reddish or yellowish red, both lighter than dark gaster; front of head shining; frontal carinae strongly divergent 21 — Body unicolored or bicolored— if bicolored then only thorax lighter than gaster; front of head opaque; frontal carinae parallel or moderately divergent dorsally 33 21(20). Scape has numerous delicate erect or suberect hairs oreas — Scape lacks erect hairs except at extreme tip, or has few scattered on inner surface near tip 22 448 Great Basin Naturalist Vol. 42, No. 4 22(21). In ventral profile, median face of clypeus on each side of keel almost flattened to form abrupt curve or angle between keel and fossa (Fig. 39) 23 — In ventral profile, upper face of clypeus not flattened, face forms a gradual curve from keel to fossa (Fig. 40) 25 23(22). Middle and hind tibiae lack erect hairs except for double row of bristles on flexor surface 24 — Middle and hind tibiae have many erect hairs in addition to double row of bristles obscuriventris 24(23). Upper surface of body lacks erect hairs; gaster pubescence thin, surface strongly shining fossaceps — Upper surface of body has abundant erect hairs; gaster pubescence dense, whitish, surface opaque laeviceps 25(22). Middle and hind tibiae have many erect hairs on all surfaces besides bristles on flexor surface 26 — Middle and hind tibiae lack or rarely have 1 or 2 erect hairs other than bristles on flexor surface 27 26(25). Head of major as broad as or broader than long; erect hairs on thorax unequal in length obscuripes — Head of major longer than broad; erect hairs on thorax short and about equal in length subnitens 27(25). Gaster densely covered with short erect hairs of plushlike appearance viewed in profile 28 — Erect hairs of gaster widely spaced, not plushlike in profile 30 28(27). Throat lacks erect hairs, or has 1 or 2 suberect ones ciliata — Throat has several to 12 or more erect hairs 29 29(28). Erect hairs on gaster short, average about 0.06 mm comata — Erect hairs on gaster long, average about 0.12 mm mucescens 30(27). Mid and hind tibiae lack row of bristles but have 3 or 4 near spur criniventris — Mid and hind tibiae have row of erect bristles that extend for one-half length or more of tibiae 31 31(30). Head of largest worker as broad as long obscuripes — Head of largest worker longer than broad 32 32(31). Throat, crest of petiole, and dorsum of thorax lack erect hairs, sometimes few present haemorrhoidalis — Throat, crest of petiole, and dorsum of thorax have numerous erect hairs, sometimes throat and petiole have only 1 or 2 integroides 33(20). Metasternum has 2 distinct hairy lobes that arise on each side of spinasternal cavity (Figs. 31, 32) 34 — Metasternum lacks such lobes 42 34(33). Head and body of imiform color 35 — Head and body each bicolored 39 35(34). Dark brown or black 36 — Pale brown or yellowish brown 38 36(35). Throat lacks erect hairs occulta — Throat has erect hairs 37 December 1982 Allred: Ants of Utah 449 37(36). Cheek and side of prothorax have erect hairs canadensis — Cheek and side of prothorax lack erect hairs altipetens 38(35). Throat lacks erect hairs neoclara — Throat has erect hairs canadensis 39(34). Throat lacks erect hairs neoclara — Throat has erect hairs 40 40(39). Yellowish brown to brownish black; gaster darker than head, which is darker than thorax; head of largest worker as broad as long; gaster pubescence thin, surface strongly shining subpolita — Gaster and upper portion of head yellowish brown to dark brown, thorax and lower part of head paler; head of largest worker longer than broad; gaster pubescence normal or dense, surface opaque or feebly shining 41 41(40). Corner of head and side of prothorax have erect hairs canadensis — Corner of head and side of prothorax lack erect hairs altipetens 42(33). Head and body of uniform color 43 — Head and body each bicolored 52 43(42). Dark brown or black 44 — Pale brown or yellowish brown 51 44(43). Cheek between eye and mandible has coarse, elongate puncta widely spaced on upper half hewitti — Cheek between eye and mandible lacks coarse, elongate puncta, or, if present, concentrated mostly on upper half 45 45(44). Throat lacks erect hairs 46 — Throat has erect hairs 50 46(45). Hairs on dorsum of 1st segment of gaster (exclusive of posterior row) usually more than 10; spinasternal cavity not surrounded by hairs (Fig. 31) 48 — Hairs on dorsum of 1st segment of gaster (exclusive of posterior- row) less than 10; spinasternal cavity surrounded by hairs 47 47(46). Length of scape greater than length of head; anterior margin of clypeus angled accreta — Length of scape less than length of head; anterior margin of clypeus broadly convex fusca 48(46). Cheek and dorsum of 1st 4 segments of gaster have dense pubescence producing a silvery luster argentea — Cheek and dorsum of 1st 4 segments of gaster have normal to thin pubescence, not silvery, but may have silky luster 49 49(48). Length of scape shorter than length of head; sides of head only slightly rounded, diverge toward base of mandibles; posterior margin of head straight or slightly convex podzolica — Length of scape equal to or longer than length of head; sides of head broadly rounded; posterior margin of head strongly convex subsericea 50(45). Length of scape greater than length of head; scale of petiole has dorsal median notch transniontanis — Length of scape not greater than length of head; scale of petiole not notched ... aetata 450 Great Basin Naturalist Vol. 42, No. 4 51(43). Throat has erect hairs aerata — Throat lacks erect hairs argentea 52(42). Throat has erect hairs 53 — Throat lacks erect hairs 54 53(52). Cheek between eye and mandible has coarse, elongate puncta, widely spaced on upper half hewitti — Cheek between eye and mandible lacks coarse, elongate puncta, or, if present, concentrated on upper half aerata 54(52). Cheek between eye and mandible has coarse, elongate puncta, widely spaced on upper half neorufibarbis — Cheek between eye and mandible lacks coarse, elongate puncta, or, if present, concentrated on upper half 55 55(54). Epinotum high with distinct angle gnava — Epinotum long and low with even convexity xerophila Key to Utah Species of Lasius Workers (Modified from Wilson 1955) 1. Eye length 0.2 times or more width of head 2 — Eye length 0.17 times or less width of head 6 2(1). Mandible has one or more offset teeth at basal angle (Fig. 5) pallitarsus — Basal tooth of mandible aligned with adjacent teeth (Fig. 6) 3 3(2). Scape lacks erect hairs; eye length usually less than 0.25 times width of head; color yellowish brown sitiens — Scape has erect hairs; eye length more than 0.25 times width of head; color other than yellowish brown 4 4(3). Next to last basal tooth of one or both mandibles markedly reduced in size rel- ative to 2 flanking teeth, or gap between next to last and last basal teeth larger than last basal tooth crypticus — Next to last basal tooth subequal in size or larger than last basal tooth, gap between them about same as last tooth 5 5(4). Scape and tibia have less than 10 or lack erect or suberect hairs alienus — Scape and tibia have more than 10 erect or suberect hairs niger 6(1). In frontal or posterior view, dorsal crest of petiole strongly convex and not notched humilis — Dorsal crest of petiole truncate or slightly convex, often notched 7 7(6). Eye has less than 35 facets 8 — Eye has 35 or more facets 9 8(6). Outer surface of tibia has numerous erect or suberect hairs fallax — Outer surface of tibia has only 1 or 2 erect or suberect hairs nearcticus 9(7). Longest hairs of posterior half of dorsum of 1st segment of gaster (exclusive of extreme posterior strip) not longer than one-half maximum width of hind tibia at midlength umbratus — Longest hairs situated as above at least three-fifths as long as tibia width 10 December 1982 Allred: Ants of Utah 451 10(9). Posterior half (except extreme posterior strip) of dorsum of 1st segment of gaster has hairs no more than semierect; erect hairs on tibia absent or sparse .... subumbratus — Posterior half (except extreme posterior strip) of dorsum of 1st segment of gaster has erect hairs; erect hairs on tibia abundant vestitus Key to Utah Species of Myrmecocystus Workers (Modified from Snelling 1976) 1. Mandible has 8 to 10 teeth; integument light yellow or brownish yellow; ocelli absent or much reduced 2 — Mandible has 6 or 7 teeth; integument not as above, but rusty red brown, black, orange or combination; ocelli well developed 5 2(1). Dorsal surface of epinotum strongly projected upward over posterior two- thirds; erect hairs sparse; upper margin of eye barely below upper margin of head pyramicus — Dorsal surface of epinotum flat or evenly convex; body often abundantly hairy; upper margin of eye well below upper margin of head 3 3(2). Head, pronotum, and gaster shiny, have few if any nonerect hairs; mid and hind tibiae have no more than 3 or 4 erect hairs beyond basal third of outer face; mesepinotal suture deeply impressed navajo — Head, pronotum, and gaster abundantly hairy; mid and hind tibiae have nu- merous erect hairs; mesepinotal suture not deeply impressed, but, if so, head length exceeds 1.3 mm 4 4(3). Mesepinotal suture impressed; epinotum as long as or longer than high, juncture of dorsal and posterior faces broadly rounded mexicanus — Mesepinotal suture not impressed; epinotum higher than long, juncture abruptly rounded or slightly angular testaceus 5(1). Uniform blackish or dark brown; anterior one-third of head may be paler; hairs on head sparse, with few erect hairs hammettensis — Bicolored or rusty red brown; head has abundant hairs, many of them erect 6 6(5). 20 or more erect hairs on cheek in front view; scape, femur, and tibia have numerous suberect hairs on all surfaces mendax — Less than 20 erect hairs on cheek; scape and femur have sparse suberect hairs 7 7(6). Cheek in frontal view has 6 or more erect hairs evenly distributed between eye and base of mandible 8 — Cheek in frontal view has no more than 4 erect hairs confined to lower half near mandible 10 8(7). Erect hairs present over at least one-half distance between inner margin of eye and base of antenna; puncta of face irregularly distributed semirufus — Erect hairs present only adjacent to inner margin of eye, do not extend more than one-fourth distance between eye and antennal base; puncta of face evenly distributed 9 9(8). Longest pronotal hairs more than one-half length of minimum diameter of eye; cheek usually has 12 to 16 erect hairs; top of head finely punctate toward sides romainei — Longest pronotal hairs less than one-half diameter of eye; cheek usually has fewer than 12 erect hairs; top of head not punctate flaviceps 452 Great Basin Naturalist Vol. 42, No. 4 10(7). Dorsum of segment 3 of gaster has dense short hairs flaviceps — Dorsum of segment 3 of gaster has few or no short hairs 11 11(10). Face has sparse short hairs; head shiny; head, thorax, and legs brownish mimicus — Face has abundant hairs; head not shiny; head, thorax, and legs rusty red brown kennedyi Acanthomyops coloradensis (Wheeler) Lasius interjectus coloradensis Wheeler, 1917, Proc. Amer. Acad. Arts, Sci. 52:532. L. claviger: Rees and Grundmann 1940:7; Cole 1942:375. A. coloradensis: Wing 1968:79; Smith 1979:1441. Records: SALT LAKE: Lake Blanche (RG). SAN JUAN: locality unknown (Wg68). UINTAH: Vernal 13.4 mi NW (A). WASATCH: Francis 14.4 mi E, Soldier Summit 3.3 mi N (A). Smith (1979:1441) lists this species from western United States, including Utah and Colorado. Most colonies have been found un- der stones. Gregg (1963:484) lists it in Colo- rado between 4654 and 8000 ft under rocks in conifers, oak, grass, and other habitats. Wing (1968) lists it from Arizona with an al- titudinal range of 2000 to 7000 ft. Three elevational records for Utah are 4750, 7100, and 7463 ft. Acanthomyops creightoni Wing Expt. A. creightoni Wing, 1968, Cornell Univ. Agric. Sta., Mem. 405:141; Smith 1979:1441. Record: GRAND: Warner Ranger Sta (Wg68). Smith (1979:1441) lists this species only from Utah. One elevational record is 9750 ft. Acanthomyops interjectus (Mayr) Lasius interjectus Mayr, 1866, Zool.-Bot. Ges. Wien, Verh. 16:888; Rees and Grundmann 1940:7; Cole 1942:375. A. interjectus: Ingham 1959:75; Wing 1968:95; Smith 1979:1441. A. claviger: Beck et al. 1967:68. Records: IRON: Cedar City (RG). JUAB: Joy (BAD). KANE: locality unknown (Wg68). SALT LAKE: Big Cottonwood Cyn (C42). Smith (1979:1441) lists this species from eastern to western United States, including Utah and Idaho, where it occurs mainly in woodlands, pastures, or meadows in exposed soil sometimes mounded, or under stones or logs. Gregg (1963:484) lists it from Colorado between 5200 and 8700 ft under rocks and logs in a variety of habitats. Wing (1968) lists it from Arizona. Cole (1942:375) indicates its habitat in Utah as under stones, one collec- tion known at 10,000 ft. Ingham (1959) found it under stones in southern Utah. Wheeler and Wheeler (1977) found it in North Dakota most frequently under rocks. Two elevational records for Utah are 5800 and 5834 ft. Beck et al. (1967:68) found it feeding on a dead pocket mouse in one instance in Utah. Acanthomyops latipes (Walsh) Formica latipes Walsh, 1862, Proc. Ent. Soc. Phila. 1:311. Lasius latipes: Rees and Grundmann 1940:7; Cole 1942:375. A. latipes: Wing 1968:101. Records: CACHE: Logan (C42). CARBON: Spring Cyn (City) (C42) ( = Storrs). DUCHESNE: Blue Bench (C42), Currant Crk (US). GARFIELD: locality unknown (Wg68). IRON: locality unknown (Wg68). SALT LAKE: locality unknown (RG). SAN JUAN: Monticello (RG). SANPETE: Majors Flats (KU). UTAH: Aspen Grove (BY). WASATCH: Currant Crk (C42). Smith (1979:1441) lists this species from eastern to western United States, including Arizona, where it occurs in open woodlands, meadows, or pastures in earthen mounds, or under stones or at the base of stumps. Gregg (1963:486) lists it from Colorado between 4800 and 8500 ft under rocks in a variety of habitats. Wing (1968) lists it from Arizona. Cole (1966:20) found its nests under stones in pinyon-juniper in southern Nevada. Wheeler and Wheeler (1963) found it frequently under rocks in North Dakota. One hundred thirty ants in three colonies in Utah were under rocks. Three collections were in sagebrush: one with matchbrush, one with snowberry, and one at the edge of aspen. Acanthomyops murphyi (Forel) Lasius murphii Forel, 1901, Ann. Soc. Ent. Belg. 45:367. A. murphiji: Wing 1968:115; Smith 1979:1441. Records: DUCHESNE: locality unknown (Wg68). SAN JUAN: locality unknown (Wg68). December 1982 Allred: Ants of Utah 453 Smith (1979:1441) designates this species as eastern to western United States, including Utah and Idaho, where it nests under stones in open woodlands. Gregg (1963:490) lists it from Colorado between 5354 and 7200 ft un- der rocks in several habitats. Wing (1968) lists it from Arizona. Acanthomyops occidentalis (Wheeler) Lasiiis occidentalis Wheeler, 1909, J. New York Ent. Soc. 17:83. A. occidentalis: Wing 1968: 124; Wheeler and Wheeler 1977; Smith 1979:1442. Record: UINTAH; locality unknown (Wg68). Smith (1979:1442) lists this species from midwestern and western United States, in- cluding Utah, Colorado, and Wyoming. Wing (1968) lists it from Arizona under stones. Aphaenogaster boulderensis M.R. Smith A. boulderensis Smith, 1941, Great Basin Nat. 2:120; Ingham 1959:47. Aecords: WASHINGTON: St. George, Zion Nat Park (159). Smith (1979:1360) lists two subspecies from the western United States, including Arizona and Nevada, probably nesting under stones. The population in Utah is likely A. h. boulderensis. Cole (1966:9) found it in south- em Nevada in mixed desert shrubs. Ingham (1963) found it in blackbrush in southern Utah. Eight elevations for Utah are between 2500 and 7066 ft. Aphaenogaster huachucana Creighton A. huachucana Creighton, 19.34, Psyche 41:189; Ingham 1959:48. Record: IRON: New Harmony (159). Smith (1979:1361) lists two subspecies from western United States, including Colo- rado and Arizona. The race in Utah is likely A. h. huachucana. Gregg (1963:340) lists it between 5100 and 6300 ft under rocks in pin- yon-juniper-oak areas in Colorado. Ingham (1959) found it under rocks in jimiper, pin- yon, sagebrush, and oak in southern Utah. Two elevation records for Utah are 5250 and 6000 ft. Aphaenogaster occidentaUs (Emery) Stenamma subterraneum occidentale Emery, 1895, Zool. Jahrb. Syst. 8:.301. A. suhterranea occidentalis: Rees and Grundmann 1940:4; Cole 1942:364; Beck et al. 1967:68. A. suhterranea valida: Creighton 1950:150; Ingham 1959:49; Beck et al. 1967:68; Smith 1979:1.362. Records (Map 1); BOX ELDER: Box Elder Cyn (US), Brigham (C42), Locomotive Spngs (BAD), Mantua Mt (C42). CACHE: Blacksmith Fk Ranger Sta (BAD), Franklin Basin (KU), Green Cyn, Hyde Park (US), Lo- gan, Logan Cyn, Providence Cyn (C42), Wellsville Cyn (US). DAVIS: Muellers Cyn (U). GRAND: Thompson (C42). IRON: Cedar City 7 mi E (A), Summit Crk (RAU). JUAB: Eureka 0.5 mi E, Nebo Loop Rd 9.9 mi S Santaquin Cyn, Ponderosa Cmpgnd (A). MILLARD: Oak Crk Cyn (US). MORGAN: Morgan (BAD). RICH: Chalk Crk (U). SALT LAKE: Big Cottonwood Cyn, But- terfield Cyn, S Dry Cyn, Ft Douglas, Holliday (C42), Little Willow Cyn (RG), Mt Olympus, Red Butte Cyn (U), Salt Lake City (RG), Univ Utah Campus (C42). SAN JUAN: Abajo Mts (U). SANPETE: Fountain Green 3 mi W (A), Majors Flats (KU). TOOELE: S Willow Cyn (U). UTAH: American Fk Cyn (U), Provo (A), N Fk Provo Cyn (U), Santaquin 3.6 and 4.6 mi E (A), Spanish Fk Cyn (KU), Thistle 2.7 and 20.4 mi E (A). WASATCH: Midway 3.7 mi NW and 5.7 mi W (A), Soapstone Cyn (U). WASHINGTON: Pine Valley (City) (BAD), Zion Nat Park (159). WEBER: Woodruff 34.8 mi W (A). Creighton (1950) and Smith (1979:1362) list two subspecies as occurring in the west- ern United States, including Utah, Colorado, and Nevada, where they nest under stones in foothill canyons. Both races have been re- corded from Utah, and may be separated as follows. Largest workers of valida are 6 mm in length and are bright red brown, whereas occidentalis workers are only 4.5 mm and are blackish brown. Gregg (1963:344) lists this species between 5354 and 7500 ft under rocks and logs predominantly in oak and de- ciduous areas in Colorado. Hunt and Snelling (1975:21) list it from Arizona. Wheeler and Wheeler (1978:391) found it between 5200 and 8400 ft in Nevada. Cole (1942:364) in- dicates its habitats in Utah as under stones and in ditch banks. Ingham (1959) lists its habitat as under stones and logs in pinyon- juniper, oak, sagebrush, and ponderosa pine in southern Utah. There were 374 ants of 11 colonies under rocks. One colony was under the same rock as a colony of Lasius pallitarsus. Seven ants were attracted to a can containing meat juices. Six colonies were in grass: one with herbs and sagebrush; one with herbs, sage- brush, and rabbitbrush; one herbs, maple, and oak; two serviceberry, maple, and oak; and one maple and oak. Two colonies were in sagebrush, one in fir, two in oak, one in fir 454 Great Basin Naturalist Vol. 42, No. 4 and oak, and one in mahogany. In 51 record- ed Utah habitats it occurred 34 times in can- yons above 5000 ft. In 27 collections where the elevation was known, it occurred about equally between 4253 and 7700 ft. One col- lection was at 9000. Eggs were found in late June and early July, and larvae in late June. In one instance, when the cover rock was removed, these ants attempted to carry the exposed larvae into their burrow. In another instance, when ants of a nearby Formica in- tegroides colony periodically crawled into the home range area of subterranea, these latter ants quickly entered their burrow. Beck et al. (1967:68) found it feeding on dead mice in four instances in Utah. Aphaenogaster uinta Wheeler A. uinta Wheeler, 1917, Proc. Amer. Acad. Arts Sci. Boston 52:517; Rees and Grundmann 1940:4; Cole 1942:364; Creighton 1950:154; Ingham 1963:77; Knowlton 1970:208, 1975:2; Smith 1979:1363. Records (Map 1): BOX ELDER: Dolphin Island (in Great Salt Lake) (C42), NW Kelton (K75) and 9 mi N (K70). DUCHESNE: NW Roosevelt (KU); EMERY: Goblin Valley 10 mi E (KU). IRON: SE Lund (163). JUAB: Chicken Crk Res (KU), Eureka 0.5 mi E (A). SALT LAKE: Big Cottonwood Cyn (U), S Dry Cyn (C42), E Mill Crk Cyn (RG), Point-of-Mt (C42), Salt Lake City (RG). Smith (1979:1363) lists this as a western in- termountain species, including Utah, Colo- rado, Nevada and Idaho, where it nests in fully exposed, dry areas. Cole (1942:364) in- dicates its nests as under stones in Utah. Ing- ham (1963) found it in alkali flats in southern Utah. Forty ants of one colony were in grass, herbs, and sagebrush under a rock. In 13 re- corded Utah habitats it occurred only 3 times in montane areas. In 7 recorded elevations between 4253 and 6425 ft, 5 were between 5000 and 5500. Brachymyrmex depilis Emery B. heeri depilis Emery, 1893, Zool. Jahrb. Syst. 7:635. B. depilis flavescens: Grundmann 1952:117. Records: BOX ELDER: WellsviUe Mts (KU). SALT LAKE: Big Cottonwood Cyn (G52). Smith (1979:1424) lists this species as mainly eastern United States, nesting under stones and wood. He does not list an inter- mountain state. Gregg (1963:449) lists it from Colorado between 4600 and 7000 ft under rocks in conifers, oak, and deciduous habitats. Wheeler and Wheeler (1963) found it fre- quently under rocks in North Dakota. Camponotus essigi M.R. Smith C. caryae var. essigi Smith, 1923, Ent. News 24:306. Records (Map 2): BOX ELDER: Hansel Mts, Mantua (KU). Smith (1979:1432) lists this species as west- ern United States, including Nevada and Idaho. Hunt and Snelling (1975:22) list it from Arizona. Wheeler and Wheeler (1978:393) found it between 6000 and 7800 ft in Nevada. One Utah elevational record is 5175 ft. Camponotus herculeanus (Linnaeus) Formica herctileana Linnaeus, 1758, Syst. Nat. Ed. 10, 1:579. C. herculeanus whijmperi: Rees and Grundmann 1940:11; Cole 1942:388; Hayward 1945:120. C. herculeanus: Smith 1979:1426. Records (Map 3): CACHE: Logan (KU), Logan Cyn (US), Monte Cristo (KU). CARBON: Scofield (BAD). DAVIS: Mueller Park (C42). DUCHESNE: Duchesne 30 mi SW (US), Mirror Lake (U), Roosevelt (C42). GAR- FIELD: Bryce Cyn Nat Park (US). JUAB: Nephi 5.5 mi W (A). RICH: Garden City, MeadowviUe (KU). SALT LAKE: Brighton (U), Cottonwood, Draper (US), Lake Blanche (C42), Red Butte Cyn (U). SAN JUAN: Mon- ticello 9.5 mi W (A). SANPETE: Bluebell Flats, Eph- raim Cyn (KU), Maple Cyn (U), Mt. Pleasant, Pleasant Crk (BAD). SUMMIT: Henrys Fk (RG), Kamas 11 mi E (U). UINTAH: Whiterocks Cyn (KU). UTAH: Aspen Grove (BY), Provo (U), Santaquin 6.7 mi E (A). WASATCH: Horse Crk (C42), Midway 11.6 mi W (A), N Fk Provo River (U). WEBER: Uintah (US). Smith (1979:1426) lists this species as east- ern to western United States, including Utah and Colorado, where it nests in rotting logs and stumps primarily in montane forest. Gregg (1963:658) lists it between 5150 and 12,500 ft under rocks and logs predominantly in conifer habitats in Colorado. Cole (1942:388) indicates its habitats as logs and stumps, particularly conifers, at high eleva- tions in Utah. One ant was taken singly in the open, three under a log, and five under a rock. Specimens were found in grass, sagebrush, and oak; chokecherry and aspen; junipers and conifers. In 33 recorded Utah habitats it oc- curred 20 times in montane forest. In 15 re- corded elevations between 4490 and 10,050 December 1982 Allred: Ants of Utah 455 ft, it occurred most frequently under 6000. When disturbed by removal of their covering these ants quickly hide. Camponotus hyatti Emery C. hyatti Emery, 1893, Zool. Jahrb. Syst. 7:669. Record (Map 2): CACHE: Green Cyn (KU). Smith (1979:1432) lists two subspecies as western United States, including Nevada, where they nest in the soil or in or under dead limbs. The Utah race is likely hyatti, which may be differentiated by its entire black or dark brown gaster, whereas in bakeri the basal two-thirds of the 1st gastric seg- ment is red. Cole (1966:19) found one nest under a juniper log in mixed brush in a desert area in southern Nevada. Camponotus laevigatus (F. Smith) Formica laevigata Smith, 1858, Cat. Hym. Brit. Mus. 6:55. C. laevigatus: Rees and Grundmann 1940:11; Cole 1942:388; Ingham 1959:70. Records (Map 2): BEAVER: Beaver Cyn (RG). CACHE: Logan, Logan Cyn (C42). GARFIELD: Car- cass Crk on Boulder Mt (U). GRAND: La Sal Mts (BY). IRON: Cedar City 7 mi E (159). SAN JUAN: Monticello 18.5 mi E (U), Navajo Mt (US). UTAH: Aspen Grove (C42), Provo (BY). WASATCH: Kamas 11 mi SE (U). WASHINGTON: Pine Valley (City) (159), Zion Nat Park (BY). WAYNE: Boulder Mt (US). Smith (1979:1426) lists the distribution of this species as western United States, includ- ing Colorado, where it nests in logs and stumps in forested areas. Hunt and Snelling (1975:22) list it from Arizona. Gregg (1963:662) lists it between 5354 and 8700 ft under logs predominantly in conifer habitats in Colorado. La Rivers (1968:7) lists it from Nevada, where Wheeler and Wheeler (1978:392) found it between 6400 and 8600 ft. Cole (1942:388) indicates its habitat as dry logs in open woods in Utah. Ingham (1959) found it in logs in ash, willow, poplar, oak, and ponderosa pine in southern Utah. In 14 recorded Utah habitats it occurred 10 times in montane forest. In 10 elevational records between 4276 and 9000 ft, it oc- curred four times under 4900, three times be- tween 7000 and 7900. Camponotus modoc Wheeler C. herculeanus var. modoc Wheeler, 1910, Ann. New York Acad. Sci. 20:299; Rees and Grundmann 1940:11; Cole 1942:388; Beck et al. 1967:68. C. pennsylvanicus modoc: Ingham 1959:69. Records (Map 3): BOX ELDER: Raft River Mts, Ta- coma Mt (U), Wellsville Mts, Willard Basin (KU). CACHE: Green Cyn, Leeds Cyn (KU), Logan, Logan Cyn (C42). CARBON: Scofield 4 and 6 mi S (A). DAG- GETT: Palisade Park (RG). DUCHESNE: Avintaquin Cmpgnd (A), Mirror Lake (BY), Trial Lake (U). EMERY: Perron Res (RG). GARFIELD: Carcass Crk on Boulder Mt (U). IRON: Cedar Breaks Nat Mon (U), Cedar City (RAU). KANE: Long Valley Jet 11 mi W (159). RICH: Garden City, Monte Cristo (KU), Woodruff 18.4 mi W (A), Wyoming brdr 0.5 mi S on U16 (A). SALT LAKE: Big Cottonwood Cyn (C42), Brighton (U), Little Willow Cyn (C42), Red Butte Cyn, Salt Lake City (U), Taylors- ville (C42). SAN JUAN: Monticello 5 mi W (U). SAN- PETE: Ephraim 8.8 mi E (A), Maple Cyn (U), Orange- ville 23.9 mi W (A). SUMMIT: Kamas 21 mi E, Mirror Lake 6.4 mi N (A), Oakley (U). TOOELE: Deep Crk Mts (U). UINTAH: Dinosaur Nat Mon (GR63), Jet Red Cloud Loop rd and U44 14 mi W (A), E Wall Lake (in Uinta Mts) (U), Whiterocks Cyn (KU). UTAH: Provo, Santaquin 6.7 and 7.7 mi E, Silver Lake Flat, Tibbie Fk Cyn (A), Utah Lake (U). WASATCH: Francis 8.1 mi E, Hanna 14.3 mi W, Soldier Summit 7.9 and 10.1 mi N (A), Strawberry Valley (C42). WASHINGTON: Kolob, Pine Valley (City) (159). COUNTY UNKNOWN: Wood- bum (C42) (no such city listed in Utah Gazetteer). Smith (1979:1426) lists this species as west- ern United States, including Colorado, where it occurs in logs and stumps in forested areas. Gregg (1963:667) lists it in Colorado between 4800 and 11,300 ft under rocks and logs pre- dominantly in conifer habitats. Hunt and Snelling (1975:22) list it from Arizona. La Rivers (1968:7) hsts it from Nevada, where Wheeler and Wheeler (1978:392) found it be- tween 6000 and 12,000 ft. They found it fre- quently in wood in North Dakota (1963). Cole (1942:388) indicates its habitat in Utah as logs and stumps of conifers at high eleva- tions. Ingham (1959) found it in southern Utah in logs in sagebrush, fir, aspen, and pon- derosa pine. Seventy-two ants in 10 collections were found under logs. In 5 of these the ants were under the same log with ants of the genus Formica: once with F. gnava, once F. podzo- lica, once F. gnava and F. neoclara, once F. fusca and F. subnuda, and once F. obscuri- ventris, F. podzolica, and F. subnitens. Thirty-two ants in 2 collections were taken from inside a log, 15 ants in 2 collections from the base of a dead standing tree, one ant under a rock, one from a dead chipmunk, 12 in 4 collections crawling in the open, and 23 of 2 collections from ground burrows. The burrows did not have mounds: one of them was in an area of Oregon grape and herbs, 456 Great Basin Naturalist Vol. 42, No. 4 where 13 ants were found, and the other, with 10 ants, was in sagebrush and grass. The burrows in the sage-grass habitat were also occupied by F. neogagates. Twelve collec- tions were in aspen: 8 in association with conifers, one with grass, herbs, and shrubs, and one grass and herbs. Four collections were in conifers; one cottonwoods; one oak, grass, herbs, and sagebrush; one herbs and Oregon grape; and one grass and sagebrush. In 54 recorded Utah habitats it occurred 42 times in montane forest. In 23 elevations be- tween 4200 and 10,399 ft, 11 collections were under 7000 and 12 over 8000. These ants are awkward crawlers, and when they are disturbed they hide under debris. Beck et al. (1957:68) found it feeding on dead rodents in three instances in Utah. Camponotus nearcticus Emery C. marginatus var. nearcticus Emery, 1893, Zool Jahrb. Syst. 7:669. C. caryae decipiens: Rees and Grundmann 1940:11. C. nearcticus decipiens: Cole 1942:388. C. nearcticus: Knowlton 1970:208, 1975:2. Records (Map 2): BOX ELDER: Kelton Pass (K70), Wildcat Hills (KU). CACHE: Logan (KU). SALT LAKE: E Mill Crk Cyn (RG). SUMMIT: Mirror Lake 17.3 mi N (A). TOOELE: Tooele (US). UINTAH: Dinosaur Nat Mon (Gr63). WASHINGTON: St George (KU). Smith (1979:1432) Usts this as eastern to western United States, including Colorado and Idaho, where it lives in trees, logs, pine cones, and other wood products. Gregg (1963:675) lists it between 4800 and 7000 ft in duff and under logs predominantly in cot- tonwood-willow habitats in Colorado. Hunt and Snelling (1975:22) list it from Arizona. Wheeler and Wheeler (1963) found it fre- quently in wood in North Dakota. Cole (1942:388) indicates its habitat in Utah as standing dead trees. One collection of 10 ants was taken from under a log in sagebrush, aspen, and conifers. In seven known specific habitats, it was taken twice in montane forest. Three known eleva- tions are between 2760 and 4923 ft. Camponotus novaeboracensis (Fitch) Formica novaeboracensis Fitch, 1845, Trans. New York State Agr. Soc. 14:766. C. herculeanus ligniperdus var. noveboracensis: Rees and Grundmann 1940:11; Cole 1942:388. C. novaeboracensis: Smith 1979:1426. Records (Map 3); CARBON: Myton rd 10.5 mi E US6 (A). DAVIS: Bountiful (U). DUCHESNE: Muellers Prk, Roosevelt (C42). GARFIELD: Carcass Crk (on Boulder Mt) (U). GRAND: Moab (U). IRON: Cedar City 7 mi E (A). RICH: Chalk Crk (U). SALT LAKE; Big Cotton- wood Cyn, Butterfield Cyn, Ft Douglas, Emigration Cyn, Red Butte Cyn, Salt Lake City (U). SAN JUAN: La Sal (U). TOOELE: S Willow Cyn (U). UTAH: American Fk Cyn (U). Smith (1979:1426) lists this species as east- ern to western United States, including Utah and Colorado, nesting in logs and stumps in wooded areas. Gregg (1963:663) lists it from Colorado between 5500 and 8500 ft under logs predominantly in conifer forest. Wheeler and Wheeler (1963) found it frequently in wood in North Dakota. One collection of four ants was from under a rock in pinyon and juniper, and the other of one ant under a log in mahogany. In 15 re- corded Utah habitats it was taken 9 times in montane areas. In 10 known elevations be- tween 4042 and 9000 ft it was found most frequently under 7000. In one collection, when the cover boulder was removed, the ex- posed ants crawled rapidly into a burrow. Camponotus ocreatus Emery C. maculatus ocreatus Emery, 1893, Zool. Jahrb. Syst. 7:673. C. ocreatus: Ingham 1959:71. Records: DUCHESNE: Duchesne 9 mi W (WU). WASHINGTON: Zion Nat Park (159). Smith (1979:1429) lists this species as southwestern United States, including Ari- zona and Nevada, nesting under stones. Cole (1966:20) found a nest under a stone in mixed shrubs in southern Nevada. Ingham (1959, 1963) found it in southern Utah under stones in sagebrush, juniper, galletagrass, rabbit- brush, winterfat, shadscale, greasewood, and alkali flats. In four recorded Utah elevations it was taken three times between 4000 and 5000 ft and once at 7000 in desert situations. Camponotus rasilis Wheeler C. fallax subsp. rasilis Wheeler, 1910, J. New York Ent. Soc. 18:227. C. rasilis: Gregg 1963:677. Record: UINTAH: Dinosaur Nat Mon (Gr63). Gregg (1963:677) lists this species from Colorado between 4800 and 6970 ft in duff and decaying logs in pinyon-juniper-oak and cottonwood-willow habitats, and shows a Utah record. Hunt and Snelling (1975:22) list it from Arizona. December 1982 Allred: Ants of Utah 457 Camponotus sansabeanus (Buckley) Formica San Sabeanus Buckley, 1866, Proc. Ent. Soc. Phila. 6:167. C. maculatus sansabeanus: Rees and Grundinann 1940:11. C. maculatus sansabeanus var. torrefactus: Rees and Grundmann 1940:11. C. sansabeanus: Cole 1942:387. C. sansabeanus torrefactus: Cole 1942:388; Creighton 1950:379; Grundmann 1958:165; Beck et al. 1967:68; Smith 1979:1429. C. sansabeanus sansabeanus: Smith 1979:1429. Records (Map 3): GRAND: Moab (G48). IRON: Paro- wan (RG). SALT LAKE: Big Cottonwood Cyn (U), Mill Crk Cyn, Salt Lake City (RG). SUMMIT: Mirror Lake 1 mi N (A). UINTAH: Dinosaur Nat Mon (Gr63). UTAH: Aspen Grove (BY). COUNTY UNKNOWN: Oris (RG) (? = Osiris in Garfield Co). Smith (1979:1429) designated three sub- species, buUmosus, sansabeanus, and torre- factus, as primarily western United States, in- cluding Utah, Colorado, Arizona, and Nevada, nesting under stones. He designated the latter two subspecies as occurring in Utah. They may be separated by the scape of the major, which is lobulate at the base in sansabeanus, but not lobulate in torrefactus. Gregg (1963:669) lists this species from Colo- rado between 4500 and 7000 ft under rocks, imder logs, and in large mounds in pinyon- jmiiper and cottonwood-willow habitats, and shows a Utah record. Cole (1942:387) in- dicates that in Utah it nests under stones in dry woods. Grundmann (1958:165) indicates that in Utah it may be found under stones in mountain brush-juniper areas. One collection of 10 ants was under a log in conifers. In eight recorded Utah habitats four were in montane forest. In seven record- ed elevations between 4042 and 10,050 ft three were under 5000. Beck et al. (1967:68) found it feeding on dead rodents in one in- stance in Utah. Camponotus semitestaceus Emery C. maculatus vicinus var. semitestaceus Emery, 1893, Zool. Jahrb. Syst. 7:668. C. semitestaceus: Allred and Cole 1979:99. Record (Map 2): KANE: Glen Cyn City (AC). Smith (1979:1429) lists this species as most- ly western United States, but no inter- mountain state is indicated. It nests under stones or in low crater mounds. Hunt and Snelling (1975:22) hst it from Arizona. Allred and Cole (1979) found it in southern Utah in ephedra-grass and sagebrush. La Rivers (1968:7) lists it from Nevada. One record in Utah was taken at 3250 ft. Camponotus vicinus Mayr C. vicinus Mayr, 1870, Verh. Zool.-Bot. Ges. Wien 20:940; Grundmann 1958:165; Ingham 1959:71; Beck et al. 1967:68; Knowhon 1970:208, 1975:2; Allred and Cole 1979:97. C. maculatus vicinus: Rees and Grundmann 1940:11. C. sansabeanus vicinus: Cole 1942:387, 388; Hayward 1945:120. C. sansabeanus vicinus var. nitidiventris: Cole 1942:387. C. sansabeanus vicinus var. luteangulus: Cole 1942:388. Records (Map 4): BEAVER: Minersville (US). BOX ELDER: Brigham, Brigham Cyn, Cedar Crk (City) (US), Dolphin Island (C42), Kelton, Kelton Pass, Locomotive Spngs (K70), Lucin (BAD), Mantua (US), Raft River Mts (BY), Snowville and 16 mi SW (KU), WiUard (C42). CACHE: Cache Jet (C42), Green Cyn (KU), Logan, Lo- gan Cyn, Providence Cyn, Trenton (C42). CARBON: Myton rd 10.5 mi E US6 (A). DAGGETT: Bridgeport (BAD) and 8 mi S (US), Browns Park (BY), Pipe Crk (US), Willow Crk (BY). DAVIS: Farmington (C42), Farmington Cyn (US). DUCHESNE: Myton (US), Roosevelt (C42), Sheep Crk (BY), Tabiona 11.6 mi E (A). EMERY: Ferron (RG). GARFIELD: Boulder Mt (G58), Bryce Cyn Nat Park (BY), Horse Valley (in Henry Mts) (U). GRAND: Dewey 3 mi N (U), Moab (RG) 14 mi N (U) and 15 mi SE (KU). IRON: Cedar City (RAU), Mod- ena 10.4 mi NE (A), Parowan (C42), Shirts Cyn (RAU). JUAB: Callao (BAD), Diamond Cyn (C42), Lynndyl Sand Dunes (BY), Nephi 12.1 mi W, Ponderosa Cmpgnd (A), Tintic, Trout Crk (City) (C42). KANE: Glen Cyn City (AC), W Mt Carmel Jet (K59), Navajo Wells (BAD), Pink Coral Sand Dunes (US). MILLARD: Candy (BAD), Swasey Spngs (RG), White Valley (C42). MORGAN: Morgan (US), Porterville, Sage Range Mts (KU). PIUTE: Fish Lake Jet 1 mi S (U), Marysvale (A). RICH: Sage Crk Jet 5.1 mi W (A). SALT LAKE: Big Cottonwood Cyn, Butterfield Cyn (C42), City Crk Cyn (U), Draper (US), Ft Douglas, Granite, E Mill Crk Cyn, Salt Lake City (C42). SAN JUAN: Abajo Mts (G58), Blanding (C42) and 8 mi N (U), Dead Horse Pt (BAD), Hole-in- the-Rock (U), La Sal Crk (RG), La Sal Jet (A), Mexican Water (BAD), Monticello 2 mi W (A) and 17 mi E (U), Nat Bridges Nat Mon (US), Navajo Mt (US), Pack Crk (US), Jet U95 and U261 7.7 mi W (A). SANPETE: Ches- ter (C42), Ephraim (US), Ephraim Cyn (U), Mt Pleasant (BAD). SEVIER: Koosharem, Paradise Valley (BAD), Sa- lina Cyn nr Fremont Jet (U). TOOELE: Benmore, Clo- ver (C42), Dolomite (US), Gransville (C42) (? = Grants- ville), Mercur (BAD), Orrs Ranch, Vernon (C42), S Willow Cyn (U), Willow Spngs (C42). UINTAH: Bo- nanza (US), Dinosaur Nat Mon (Gr63), Dry Fk rd 13.4 mi N U121 (A), Duchesne 9 mi W (WU) (? = Ft Du- chesne), Pelican Lake (US). UTAH: American Fk (RG), Jordan Narrows, Mercur (US), Prove, Springville, Thistle (BY), Tibbie Fk Cyn, Tibbie Fk Lake 0.5 mi W (A). WASATCH: Deer Crk Res (U), Hanna 9.2 mi W, Mid- way 3.7 mi NW (A). WASHINGTON: Pine Valley (City), nr Short Crk (Arizona) (159), Sunset Cyn (nr Vir- gin) (RAU), Zion Nat Park (BY). WAYNE: Fruita 5 mi SE (U), Torrey (BAD). WEBER: Ogden (C42). 458 Great Basin Naturalist Vol. 42, No. 4 Smith (1979:1430) lists this species from western United States, including Colorado, nesting under stones or rotting wood in soil. Gregg (1963:674) Hsts it between 3500 and 9600 ft. under rocks and logs in a variety of habitats in Colorado. Hunt and Snelling (1975:22) list it from Arizona. Cole (1966:20) found it in southern Nevada under rocks commonly in pinyon-juniper, also in open dry areas in Utah (1942:387). Wheeler and Wheeler (1978:393) found it between 6000 and 9100 ft in Nevada, and frequently under rocks in North Dakota (1963). It occurs in Utah under stones in juniper-brush habitats (Grundmann 1958:165). Ingham (1959, 1963) found it under rocks in pinyon-juniper, sage- brush, serviceberry, bur sage, rabbitbrush, galletagrass and greasewood in southern Utah. Allred and Cole (1979:97) found it in southern Utah in habitats of ephedra-van- clevea-grass, juniper-ephedra-grass, sage- brush, juniper-pinyon, grass-sagebrush, and mostly in blackbrush. They also found it in abundance in associations of sagebrush-grass, sagebrush-grass-rabbitbrush, and sagebrush- rabbitbrush and less abundant in a variety of other vegetative types in Idaho (Allred and Cole 1971:239). There were 186 ants in 12 collections found under rocks, 3 in one collection singly in an open area, and 2 in one collection un- der a juniper log. In a juniper-sagebrush area in late June 12 miles west of Nephi in Juab County, 20 ants in one collection were found in a mound of Pogonomyrmex occidentalis, al- though no harvester ants were present. The carpenter ants had excavated a large cham- ber (or cleared out and enlarged the food cache area) about eight inches from the top of the mound. About 50 workers and 50 pupae were present. Eggs were found imder one rock in late June. When the rock was re- moved, the ants attempted to carry the ex- posed eggs into the burrow. Ten collections were in sagebrush: one in association with grass and herbs, one grass and juniper, one pinyon and juniper, one pin- yon, one snowberry and oak, and one snow- berry. One collection was in pinyon and juni- per; one grass and oak; one oak; one grass, herbs, maple and oak; and two aspen and fir. In 124 recorded Utah habitats 35 were in montane forest. In 69 recorded elevations be- tween 3250 and 9000 ft, 51 collections were between 4000 and 6000. Only one was under 4000, two over 8000. Beck et al. (1967:68) found it feeding on dead rodents in 13 in- stances in Utah. Conomyrma bicolor (Wheeler) Dorymyrmex pyramicus var. bicolor Wheeler, 1906, Bull. Amer. Mus. Nat. Hist. 22:342; Cole 1942:372; Grundmann 1958:164; Ingham 1959:64. D. bicolor: Beck et al. 1967:69. C. bicolor: Allred and Cole 1979:99; Smith 1979:1420. Records (Map 5): CACHE: Hodges Cyn (KU). EMERY: Greenriver (U). GRAND: Dewey (U), Crescent Jet 5 and 18.8 mi S (A), Moab (C42). IRON: Cedar City 6.3 mi W (A), Columbia Iron Mines 7 mi W, Kanaraville (159), Modena 5.3 mi W (A). JUAB: Callao (BAD), Provo 30 mi S (C42). KANE: Adairville (BAD), Kanab (C42), 5 mi E (A) and 5 mi N (159). MILLARD: Desert Range Exp Sta (U). SAN JUAN: Blanding, Bluff, Glen Cyn Res, Hole-in-the-Rock (G58), La Sal Jet (A), Lost Eden, Mexi- can Hat (G.58), Monticello (US), Nat Bridges Nat Mon (U), Rock Crk (G58). UINTAH: Bonanza 15 mi N, Jen- sen 7.5 mi E (A), Vernal (US). WASHINGTON: Grafton, Harrisburg Jet toward Hurricane, LaVerkin, Pintura, Rockville, Shivwits Indian Res, nr Shortcreek (Arizona), Snow Cyn, Springdale, Toquerville, Virgin City, Zion Nat Park (159). WAYNE: Hanksville 17 mi S (US). Smith (1979:1420) lists this species from southwestern United States, including Utah, Arizona, and Nevada, where it nests in ir- regular or craterlike mounds in exposed situa- tions. Cole (1966:18) found it commonly in craterlike mounds in creosote bush habitats in southern Nevada. In Utah it inhabits lower elevations along canyon bottoms, where it builds craterlike mounds (Grundmann 1958:164). Ingham (1959, 1963) found it in southern Utah in flat or low crater mounds in creosote bush, sagebrush, pinyon-juniper, ponderosa pine, blackbrush, joshua trees, rab- bitbrush, bur sage, sand sagebrush, four-wing saltbush, and shadscale. Allred and Cole (1979:99) found it in blackbrush in southern Utah. There were 125 ants in four collections taken from small crater mounds less than one-half inch in height and about six inches in diameter, with the dorsal opening in the crater. One mound was at the edge of a clearing of an active colony of Pogonomyr- mex occidentalis. One collection of 30 ants was taken from a burrow with no mound, in which winged forms were more abundant than the workers. Thirty ants in one collec- tion were taken from a safari; the ants scur- ried out of line when disturbed. Twenty-two December 1982 Allred: Ants of Utah 459 ants were taken in two collections crawling singly away from their burrows. These ants are fast runners. Occasionally many mounds may occur in close vicinity. Six collections were in grass: two in association with sage- brush, one old man sage, one greasewood, and one juniper. One collection was in sage- brush and matchbrush. Only once in 43 re- corded Utah localities was it found in a mon- tane area. It was taken between 2500 and 7066 ft at 35 known elevations, 24 of these between 4000 and 5000 ft. It was taken over 6000 ft only twice. In one instance a nest of Formica obtusopi- losa, around which bicolor workers were scurrying, was excavated five miles west of Modena in Iron County. When the eggs of the Formica were exposed, the bicolor ants ran into the excavated area and began car- rying the eggs away. In the same area two harvester ants of Pogonomyrmex occidentalis were "tending" a bicolor, apparently dead, at the fringe of the harvester mound. Several bicolor ants kept darting at the harvester ants, not actually making contact, in what appeared to be a type of rescue effort. The harvesters demonstrated no apparent re- sponse to the aggressive bicolor ants. Beck et al. (1967:69) found it feeding on dead rodents in two instances in Utah. Conomynna insana (Buckley) Formica insana Buckley, 1866, Proc. Ent. Soc. Phila. 6:165. Dorymyrmex pyramicus: Rees and Grundmann 1940:6; Cole 1942:371; Creighton 1950:348; Ingham 1959:64; Beck et al. 1967:69. D. pyramicus flavus: Cole 1942:371. D. pyramicus pyramicus: Grundmann 1958: 164. C. insana: Allred and Cole 1979:97. Records (Map 5): BEAVER: Frisco (BAD). BOX EL- DER: Snowville (KU). CACHE: Leeds Cyn (KU). CAR- BON: Price 20 mi E (WU). DAVIS: Layton 1.5 mi S (A). DUCHESNE: Blue Bench (C42), Duchesne (BAD), My- ton (US), Roosevelt (BAD). EMERY: Goblin Valley (BAD) and 10 mi E (KU), Greenriver (C42), Hanksville 16 mi N (KU), Hideout Cyn nr Green River (U), Hun- tington (BAD). GARFIELD: Escalante 20 mi E (U), Hite (BAD). GRAND: Arches Nat Park (WU), Dewey (U), Moab (C42). IRON: Cedar City (159), 3.7 mi E and 16.8 mi W (A), Little Pinto (US), Lund 19 mi N, Modena 10.4 mi NE (A), New Harmony 3 mi NW (159). JUAB: Nephi 5.5 mi W (A). KANE: Glen Cyn City (AC), Kanab (RG), Wahweap (KU). MILLARD: Desert Range Exp Sta (BAD), White Valley (C42). PIUTE: Kingston (BAD). SALT LAKE: Butterfield Cyn, Granite (C42), Mt Olympus (U), Parleys Cyn (C42), Riverton 2.6 mi S (A). SAN JUAN: Aztec Cyn (U), Blanding (G58), Bluff (KU) and 11 mi N (U), Hole-in-the-Rock Cyn (U), La Sal Jet 1.6 and 23.1 mi S (A), Monticello (G58), Nat Bridges Nat Mon (U), Squaw Flat Cmpgnd (in Canyonlands Nat Park) (WU), Jet U95 and U261 7.7 mi W, Jet U261 and US163 19.6 mi N (A). SANPETE: Yuba Res (BAD). TOOELE: Clive, Clover, Orrs Ranch, Tooele (C42). UINTAH: Bonanza (KU), Dinosaur Nat Mon (BAD), Ft Duchesne, Gusher (C42), Jensen (U) and 5.5 mi W (A), Lapoint, Ouray Valley (C42), Vernal 10 mi N (A). UTAH: Goshen, Lehi, Orem, Provo (C42). WASHING- TON: Central (159), Diamond Valley (BAD), Hurricane (C42), Leeds (US), Leeds Cyn (KU), Pine Valley Res (159), Rockville (BAD), Snow Cyn (KU), St George (C42), Toquerville (BAD), Virgin (KU), Zion Nat Park (WU). WAYNE: Capitol Reef Nat Park (WU), Fruita, Hanks- ville (BAD), and 17 mi S (KU), Loa (A), Torrey (BAD). Smith (1979:1420) lists this species as east- ern to western United States (no inter- motintain state listed), where it nests in ir- regular or craterlike mounds in open areas. Gregg (1963:434) lists it from Colorado be- tween 3500 and 8500 ft under rocks in a vari- ety of habitats, predominantly pinyon-juniper and grass. Hunt and Snelling (1975:22) list it from Arizona. Wheeler and Wheeler (1963) found it frequently in soil craters in North Dakota. Cole (1966:18) found it commonly in creosote bush habitats in southern Nevada. It forms craterlike nests in Utah (Cole 1942:371). Ingham (1959, 1963) found it in southern Utah in pinyon-juniper, sagebrush, creosote bush, mesquite, galletagrass, rabbit- brush, winterfat, shadscale, greasewood, and alkali flats. It was associated with Solenopsis molesta in one nest.. Allred and Cole (1979:97) found it in a large variety of desert shrub associations in southern Utah, com- monly in juniper-pinyon. There were 178 ants of seven collections taken from small crater mounds. Two were in the cleared area of active colonies of Pogono- myrmex occidentalis. In Davis County 1.5 mi south of Layton on highway US89, many small craterlike mounds were in close vicin- ity and surrounding a mound occupied by oc- cidentalis. Ten ants in one collection were taken from a mound actually on an occupied mound of occidentalis. Another mound occu- pied by occidentalis was covered with sticks, and many (30 collected) insana were crawl- ing among the sticks. No openings or mounds for these small ants were seen, nor were any of their mounds in the vicinity of the har- vester mound. No evidence was seen of the harvesters attacking the insana, and the two 460 Great Basin Naturalist Vol. 42, No. 4 species seemingly intermingled in their movements over the large mound. Nineteen miles north of Lund on the road to Pine Val- ley in Iron County, nine occupied mounds were found on a large flattened mound of oc- cidentalis. In San Juan County, 19.6 mi north of the junction of US163 and U261 along U261, insana had parts of three occidentalis with some appendages and parts of the body missing, as though chewed by the insana. Eighty ants in two collections were taken from safaris; in one case some of the ants were grouped around a dead moth. Forty- two ants in two collections were taken under rocks. In one of these colonies eggs were present in late Jime. Eight collections were in sagebrush: two in association with grass, one clover, one rabbitbrush, one juniper, one pin- yon, and one jimiper and pinyon. One collec- tion was in grass and clover, one grass and herbs, one herbs, one matchbrush and grease- wood, one rabbitbrush, and one juniper. In 87 recorded Utah localities it occurred only 5 times in montane areas. In 53 recorded eleva- tions between 2750 and 7066 ft it was taken most frequently (32 times) between 4000 and 6000. Beck et al. (1967:69) found it feeding on dead rodents in 18 instances in Utah. Crematogaster cerasi (Fitch) Myrmica cerasi Fitch, 1855 (1854), Trans. New York State Agr. Soc. 14:835. Record: WASHINGTON: Leeds (US). Smith (1979:1378) hsts this species as east- em to midwestem United States, no inter- mountain state listed, where it nests under rocks and logs. The one collection in Utah re- portedly taken at 2750 ft is questionable. Crematogaster coarctata Mayr C. coarctata Mayr, 1870, Verb. Zool.-Bot. Ges. Wien 20:990. C. vermiculata: Gole 1942:363. Record: JUAB: Provo 30 mi S (C42). Smith (1979:1378) hsts this species as west- em United States, including Nevada, nesting under rocks. Cole (1966:16) found its nests in southern Nevada in open soil and under stones in pinyon-juniper and a variety of desert shrub types; it also occurs in sagebrush in Utah (Cole 1942:363). One Utah collection was taken at 4909 ft. Crematogaster depilis Wheeler C. lineolata opaca var. depilis Wheeler, 1908, Bull. Amer. Mus. Nat. Hist. 24:478. C. depilis: Beck et al. 1967:68; Allred and Cole 1979:97. Records (Map 6): KANE: Glen Cyn City (AC). WASHINGTON: Beaver Dam Wash, Toquerville (BAD). Smith (1979:1378) lists this species as west- ern United States, including Arizona and Ne- vada, nesting in roots and lower stems of desert plants. Cole (1966:16) found it in open areas in southern Nevada in various desert shrub types, none in pinyon-juniper. Allred and Cole (1979:97) found it in a variety of desert shrub habitats, most frequently in eph- edra-vanclevea-grass and juniper-ephedra- grass associations in southern Utah. In three desert collections in Utah it was taken betwen 2350 and 3371 ft. Beck et al. (1967:68) found it feeding on dead rodents in five instances in Utah. Crematogaster emeryana Creighton C. lineolata emeryana Creighton, 1950, Bull. Mus. Comp. Zool. 104:213; Grundmann 1958:163; Beck et al. 1967:68. C. lineolata nr cerasi: Rees and Grundmann 1940:4; Cole 1942:363. C. lineolata: Ingham 1959:55. C. punctulata: Beck et al. 1967:69. C. emeryana: Smith 1979:1378. Records (Map 6): BEAVER: Minersville (BAD). BOX ELDER: Locomotive Spngs (BAD), Park Valley (City) (C42). CACHE: Logan (C42). JUAB: Callao (BAD). KANE: Mt. Carmel Jet (159), Navajo Wells (BAD). MIL- LARD: Swasey Spngs (RG). SALT LAKE: Parleys Cyn (U). SAN JUAN: Blanding, Bluff, Monticello (G58), Montezuma Crk (BAD). TOOELE: Clover, Delle, Fish- er Pass, Low (C42). UTAH: American Fk Cyn (U), Chimney Rock Pass (BAD), Provo (C42). WASHING- TON: Diamond Valley (BAD), Grafton, La Verkin, New Harmony, Pintura, Rockville, Santa Clara River, Zion Nat Park (159). WEBER: Ogden Cyn (U). Smith (1979:1378) lists this species as west- ern United States, including Utah, Colorado, and Arizona, where it usually nests under rocks in mountains at elevations over 6000 ft. Gregg (1963) lists it between 3500 and 8000 ft in a variety of habitats, predominantly un- der rocks and logs in oak and pinyon-juniper areas in Colorado. Cole (1942:363) indicates its habitats in Utah as under stones, logs, and bark. Ingham (1959, 1963) found it under stones and wood in willow, poplar, tamarix, oak, juniper, ash, sagebrush, squawbrush, and shadscale in southern Utah. December 1982 Allred: Ants of Utah 461 In 29 localities, only three were in mon- tane forest. In 25 known elevations between 3000 and 7066 ft, 10 were between 4000 and 5000. Beck et al. (1967:68) found it feeding on dead rodents in eight instances in Utah. Crematogaster hespera Buren C. hespera Buren, 1968, J. Georgia Ent. Soc. 3:98; Smith 1979:1379. Record: WASHINGTON: Zion Nat Park (B). Smith (1979:1379) lists this species as west- em United States, including Utah and Ari- zona, principally as an arboreal form in cot- tonwood trees. Buren (1968:100) indicates that it is largely an arboreal species of cot- tonwoods and other trees, but also occurs in logs at elevations between 2000 and 5500 ft. One Utah collection was taken at 4276 ft. Crematogaster minutissima Emery C. victima missouriensis Emerv, 1895, Zool. Jahrb. Syst. 8:287. C. minutissima: Beck et al. 1967:69. Record: WASHINGTON: Diamond Valley (BAD). Creighton (1950) and Smith (1979) list three subspecies as eastern to western United States, including Colorado and Arizona, nest- ing in soil at the base of stumps. The Utah form is likely smithi, although missouriensis may come into eastern Utah. The two may be separated by the smooth shining dorsum of the promesonotum in smithi, whereas in missouriensis it is finely punctate. Gregg (1963:364) lists this species at 4150 ft under rocks in grassy habitats in Colorado. Beck et al. (1967:69) found it feeding on dead rodents in one instance in Utah. Crematogaster mormonum Emery C. lineolata coarctata var. mormonum Emery, 1895, Zool. Jahrb. Syst. 8:284; Rees and Grundmann 1940:4. C. coarctata var. mormonum: Cole 1942:363. C. mormonum: Creighton 1950:215; Ingham 1959:56; Beck et al. 1967:69; Knowlton 1970:209; 1975:2; Allred and Cole 1979:99; Smith 1979:1380. Records (Map 6): BEAVER: Frisco (BAD). BOX EL- DER: Hansel Mts (K75), Snowville (KU) and 4 mi W (K70). CACHE: Leeds Cyn (KU). JUAB: Joy (BAD). KANE: Glen Cyn City (AC), Kanab (US). MILLARD: Swasey Spngs (BAD). SALT LAKE: E Mill Crk Cyn, Salt Lake City (RG). SAN JUAN: Bluff (KU). SEVIER: Koosharem (BAD). TOOELE: Stansburv Island (C42). WASHINGTON: Leeds (KU), Pine Valley Res (159), Zion Nat Park (KU). Smith (1979:1380) hsts this species as west- ern United States, including Utah, Nevada, and Idaho, nesting under rocks. Cole (1942:363) indicates its habitats in Utah as under stones and logs. Ingham (1959) found it under stones and logs in southern Utah. All- red and Cole (1979:99) found it in a saltbush- sagebrush association in southern Utah. In 17 recorded Utah habitats it was taken only 3 times in montane forest. In 12 re- corded elevations between 2750 and 6850 ft it was taken 6 times between 4000 and 5000, 4 times above 6000. Beck et al. (1967:69) found it feeding on dead rodents in four in- stances in Utah. Crematogaster noctuma Buren C. noctuma Buren, 1968, J. Georgia Ent. Soc. 3:112; Knowlton 1975:2; Smith 1979:1380. Records: BOX ELDER: Hansel Mts (K75). SAN JUAN: Nat Bridges Nat Mon (B). Smith (1979:1380) lists this species as west- ern United States, including Utah and Ari- zona. In one Utah collection it was taken at 5700 ft. Forelius foetidus (Buckley) Formica foetida Buckley, 1866, Ent. Soc. Phila., Proc. 6:167. Forelius foetida: Ingham 1959:64. Records: EMERY: San Rafael Swell (U). JUAB: Chicken Crk Res (KU). KANE: Grosvenor Arch (US). WASHINGTON: Rockville (159), Smith (1979:1419) hsts this species as mid- west to western United States, including Col- orado, nesting under various objects or in small craters. Gregg (1963:434) lists it be- tween 4400 and 4900 ft under rocks in cot- tonwood-willow and grass habitats in Colo- rado. Hunt and Snelling (1975:22) list it from Arizona. LaRivers (1968:7) hsts it from Ne- vada. Ingham (1959) found it in small crater mounds in sagebrush, blackbrush, cliffrose, Russian thistle, and cheatgrass in southern Utah. In four localities in Utah it was taken between 3800 and 5000 ft in desert areas. Formica accreta Francoeur F. accreta Francoeur, 1973, Soc. Ent. du Quebec, Mem. no. 3, pp 182-9. Record: WEBER: Beaver Crk (KU). Smith (1979:1452) lists this species from western United States, including Idaho. 462 Great Basin Naturalist Vol. 42, No. 4 Formica aetata Francoeur F. aerata Francoeur, 1973, Soc. Ent. du Quebec, Mem. no. 3, pp. 116-22. Record: CACHE: Tony Grove Lake (KU). Smith (1979:1452) lists this species from western United States, including Nevada, nesting mider rocks. In one Utah collection it was taken at 8075 ft. Formica altipetens Wheeler F. cinerea cinerea var. altipetens Wheeler, 1913, Bull. Mus. Comp. Zool. 53:399. F. cinerea altipetens: Rees and Grundmann 1940:10; Cole 1942:383. F. altipetens: Creighton 1950:531; Francoeur 1973:58; Smith 1979:1452. Records (Map 7): CACHE: Ant Valley, Antelope Val- ley, Blacksmith Fk Cyn, Elk Valley, Franklin Basin, Lo- gan Cyn, Petersboro, Pole Crk Spng, Rock Crk, Tony Grove (KU), Wellsville Cyn (F). DUCHESNE: Fruitland (U). GARFIELD: Boulder Mt (U). GRAND: Warner Ranger Sta (RG). RICH: Garden City (KU). SAN JUAN: Monticello (KU). SUMMIT: Henrys Fk Basin (RG). TOOELE: S Willow Cyn (U). WASATCH: Soldier Sum- mit 1 mi E (A). Smith (1979:1452) lists this species from western United States, including Utah, Colo- rado, Arizona, Idaho, and Wyoming, nesting in open areas sometimes with low mounds or under objects. Gregg (1963:507) lists it be- tween 6000 and 11,000 ft under rocks and logs mostly in conifer areas in Colorado. La Rivers (1968:9) hsts it from Nevada. Wheeler and Wheeler (1977) found it frequently in thatched or earthen mounds and under rocks in North Dakota. Cole (1942:383) indicates its habitats in Utah as under stones or in flat earthen mounds in open areas. Four ants in one collection were found crawling singly on the ground in sagebrush and herbs. In 19 recorded Utah habitats 15 were in montane forest. In nine recorded ele- vations between 4471 and 10,500 ft it was found most frequently (four times) between 7000 and 9000. Formica argentea Wheeler F. fusca var. argentata Wheeler, 1902, Amer. Nat. 36:952; Cole 1942:383. F. fusca argentea: Rees and Grundmann 1940:9; Knowl- ton 1970:209. F. argentea: Francoeur 1973:150. Records (Map 7): BEAVER: Beaver Crk (F). BOX EL- DER: Cedar Hill, Clear Crk Cyn (KU), Kelton Pass (K70), SnowviUe (C42), Willard Basin (in Wellsville Mts) (KU). CACHE: Ant Valley, Bear River Mts (KU), Black- smith Fk Cyn (C42), Elk Valley, Franklin Basin, High Crk, Leeds Cyn, Logan (KU), Logan Cyn (C42), Mendon (F), Paradise, Petersboro, Providence, Tony Grove Cyn, Tony Grove Lake (KU), Wellsville (F). DAGGETT: Sheep Crk (RG). DAVIS: Farmington, Farmington Cyn (KU). EMERY: Hideout Cyn nr Green River (U). GAR- FIELD: Orton (KU). JUAB: Ponderosa Cmpgnd, Red Crk Spng (KU). KANE: Mt Carmel (F), Wahweep (KU). RICH: Allen Cyn, Monte Cristo and 6 mi S, Randolph (WU). SALT LAKE: Garfield (F), Little Willow Cyn (C42). SAN JUAN: Monticello (RG) and 2 mi W (A). SE- VIER: Burrville (U), Salina Cyn nr Fremont Jet (U). TOOELE: Delle (C42). UINTAH: Jet Dry Fk rd and Red Cloud Loop rd. Jet Red Cloud Loop rd and U44 9.2 mi W (A), Whiterocks (KU). UTAH: American Fk (RG), Spanish Fk Cyn (KU). WASATCH: Midway 3.7 mi NW, Strawberry Res 4 mi S (A). WASHINGTON: Pine Valley (City) (KU). WAYNE: Capitol Reef Nat Park (U). WE- BER: head of Beaver Crk, Monte Cristo (KU). COUNTY UNKNOWN: Rand (F) (? = abbr for Randolph in Rich Co). Smith (1979:1452) lists this species from eastern to western United States including Arizona, nesting in open or semiopen areas under rocks or in low mounds. Gregg (1963:523) hsts it from Colorado between 4500 and 11,000 ft under rocks and wood in a variety of habitats. La Rivers (1968:9) lists it from Nevada, where Wheeler and Wheeler (1978:394) found it between 6200 and 11,500 ft. They also found it frequently imder rocks, also in earthen mounds in North Dakota (1977). Cole (1942:383) indicates its habitats in Utah as under stones and logs in cold for- ests at higher elevations. Twenty-four ants in four collections were taken from under rocks. In one collection it was under the same rock with Formica la- sioides, once F. fusca, and once F. pallide- fulva and Pheidole desertorum. Four speci- mens in one collection were taken singly crawling in the open. Two collections were in oak, one associated with grass. One collec- tion was in legumes and sagebrush, one a grassy meadow, and one grass and sagebrush in an open area of aspen and pine. In 54 re- corded Utah habitats it was taken 28 times in montane forest. In 23 recorded elevations be- tween 4225 and 9300 ft it was taken 10 times under 5000, 12 times between 6000 and 9000. Formica calviceps Cole F. calviceps Cole, 1954, J. Tenn. Acad. Sci. 29:164. Record: UTAH: Spanish Cyn (KU) (? = Spanish Fk Cyn). December 1982 Allred: Ants of Utah 463 Smith (1979:1457) lists this species only from New Mexico nesting under stones banked with detritus. Its occurrence in Utah is questionable. Formica canadensis Santschi F. cinerea var. canadensis Santschi, 1913, Ann. Soc. Ent. Belg. 57:435; Knowlton 1975:2. F. cinerea: Rees and Grundmann 1940:10; Knowlton 1970:209. F. cinerea neocinerea: Rees and Grundmann 1940:10; Gole 1942:383; Knowlton 1970:209, 1975:2. F. cinerea lepida: Creighton 1950:531; Beck et al. 1967:69. F. montana: Ingham 1959:82. F. canadensis: Francoeur 1973:66; Smith 1979:1452. F. cinerea canadensis: Knowlton 1975:2. Records (Map 7): BEAVER: Puffers Woke (F) (? = Puffer Lake). BOX ELDER: Locomotive Spngs (K70), Park Valley (City) (KU), Snowville (C42), Wellsville Mts, Willard Basin (KU). CACHE: near Franklin (Idaho), Franklin Basin, Hyrum, Logan Cyn, Tony Grove Lake (KU), Wellsville (RG). IRON: Cedar Breaks Nat Mon, Cedar City (F), Newcastle (159). KANE: Kanab (C42), Navajo Lake (159). RICH: Laketown, Randolph (KU). SALT LAKE: Alta (U), Wasatch (RG). SAN JUAN: La Sal Mts (U), Montezuma Crk (BAD). SANPETE: Ephra- im Cyn (KU). SEVIER: Venice (F). SUMMIT: Wasatch (RG). TOOELE: Clover (F). UINTAH: Bonanza (KU), Ft Duchesne (F), Jensen 17 mi SW (C42), Lapoint (F), Neola 2.7 mi E (A), Paradise Park (U), Vernal, White- rocks (F). UTAH: Goshen (F), Lehi (BAD). WEBER: Beaver Crk (KU). Smith (1979:1452) lists this species from midwest to western United States, including Utah, Colorado, Arizona, Idaho, and Wyo- ming, nesting in open or wooded areas in soil, sometimes with a low mound. Gregg (1963:511) lists it from Colorado between 3500 and 11,000 ft under rocks, wood, and in thatched hummocks predominantly in mead- ow situations. Allred and Cole (1971:239) found it rarely in associations of sagebrush- grass in Idaho. La Rivers (1968:9) lists it from Nevada. Wheeler and Wheeler (1977) found it frequently in earthen mounds, also com- monly under rocks in North Dakota. Cole (1942:383) indicates its habitats as under stones or flat earthen mounds in open areas in Utah. Ingham (1959) found it in southern Utah under stones in sagebrush, aspen, fir, and spruce. Knowlton (1975:2) found it asso- ciated with rabbitbrush in northern Utah. Forty ants in one collection were taken from the ground in grass near an irrigated field. In 38 recorded Utah habitats 13 were in montane forest. In 29 recorded elevations between 4495 and 10,399 ft, 18 were under 6000 and 7 were over 9000. Formica ciliata Mayr F. ciliata Mayr, 1886, Verh. Zool.-Bot. Ges. Wien 36:428; Rees and Grundmann 1940:9; Cole 1942:379; Creighton 1950:483; Grundmann 1958:165; Gregg 1963:546; Smith 1979:1457. Records: GRAND: Warner Ranger Sta (Gr63). SAN JUAN: Blanding (RG), between Blanding and Vedura (C42) (? = Verdure). SANPETE; Wales 3.3 mi W (A). Smith (1979:1457) lists this as a species of western United States, including Utah, Colo- rado, and Wyoming, with nests (sometimes thatched) in meadows or open woods. Gregg (1963:546) lists it between 5354 and 11,000 ft under rocks and logs principally in conifer habitats in Colorado, and lists a record for Utah. Wheeler and Wheeler (1963) found it only under rocks in North Dakota. Cole (1942:379) indicates its habitats as under stones or logs in Utah. Nests of this species are usually low, thatched mounds in mead- ows and open woods (Grvmdmann 1958:165). Seven ants in one collection in Utah were taken from under a rock in sagebrush and rabbitbrush near a meadow. One recorded elevation in Utah is 6103 ft. Formica comata Wheeler F. comata Wheeler, 1909, J. New York Ent. Soc. 17:85; Rees and Grundmann 1940:9; Cole 1942:379. Records: GARFIELD: Carcass Crk (on Boulder Mt) (U). SALT LAKE: Mill Crk Cyn (RG). SAN JUAN: La Sal Mts (U). Smith (1979:1457) lists this species as west- ern United States, including Colorado, nest- ing under stones banked with thatch. Gregg (1963:550) lists it between 6000 and 7704 ft in sagebrush, chaparral, and grass habitats in Colorado. Cole (1942:379) indicates its habi- tats in Utah as under stones or logs, generally banked or covered with detritus. Two Utah collections were taken at 9000 ft. Formica criniventris Wheeler F. crinita Wheeler, 1909, J. New York Ent. Soc. 17:87. F. crinoventris: Rees and Grundmann 1940:9. F. criniventris: Cole 1942:379; Beck et al. 1967:69; Smith 1979:1457. Records (Map 8): BOX ELDER: Garland (KU). CACHE: Blacksmith Fk Cyn (KU). SALT LAKE: Big Cottonwood Cyn, Brighton (U). SEVIER: Paradise Val- ley (BAD). COUNTY UNKNOWN: Boulton (RG) (? = Boulter in Juab Co). 464 Great Basin Naturalist Vol. 42, No. 4 Smith (1979:1457) lists this species as west- em United States, including Utah and Colo- rado, nesting under stones banked with thatch in meadows and open forests. Gregg (1963:550) lists it between 5100 and 5900 ft under rocks predominantly in gardens and open meadows in Colorado. Wheeler and Wheeler (1963) found it frequently under rocks in North Dakota. Cole (1942:379) in- dicates its habitat in Utah as under stones banded with detritus. Knowlton (1975:2) found it associated with rabbitbrush in north- em Utah. In six recorded Utah habitats it was taken four times in montane forest. Three known elevations are 4340, 5000, and 8600 ft. Beck et al. (1967:69) found it feeding on dead ro- dents in one instance in Utah. Formica densiventris Viereck F. fmca var. densiventris Viereck, 1903, Trans. Amer. Ent. Soc. 29;74. F. rasilis spicata: Creighton 1950:507. F. densiventris: Smith 1979:1462. Records (Map 8): BOX ELDER: Snowville (KU). CACHE: Bear River Mts, Leeds Cyn (KU). RICH: Bear River, Wasatch Mts (KU). Smith (1979:1462) hsts this species from western United States, including Utah and Colorado, nesting in forests under stones and logs, occasionally thatched. Hunt and Snell- ing (1975:23) list it from Arizona. Wheeler and Wheeler (1978:394) found it between 7200 and 10,000 ft in Nevada. One collection in Utah was taken at 4544 ft. Formica emeryi Wheeler F. emeryi Wheeler, 1913, Bull. Mus. Comp. Zool. 53:389. Record: CACHE: Cowley Cyn (KU). Smith (1979:1464) lists this species only from Colorado, where it associates with F. neogagates. Gregg (1963:606) lists it at 6000 ft in meadow habitats in Colorado. Formica fossaceps Buren F. fossaceps Buren, 1942, Iowa State Coll. J. Sci. 16:402. Record: BOX ELDER: Tremonton (KU). Smith (1979:1458) lists this species from midwestem United States (no intermountain state listed), where it nests under stones or logs banked with thatch or in low earthen mounds covered with thatch or grass. Wheel- er and Wheeler (1963) found it under rocks and in earthen mounds in North Dakota. One collection in Utah at 4315 ft is questionable. Formica fusca Linnaeus F. fusca Linnaeus, 1758, Syst. Nat. (10 ed.) 1:580; Cole 1942:381, 382; Creighton 1950:532; Grundmann 1958:165; Ingham 1959:81; Beck et al. 1967:69; Knowlton 1970:209, 1975:2. F. fusca subaenescens: Rees and Grundmann 1940:9; Cole 1942:382. F. fusca subsericea: Cole 1942:382. F. marcida: Creighton 1950:534; Ingham 1959:82. Records (Map 8): BEAVER: Beaver 5.5 mi E (U). BOX ELDER: Brigham (C42), Cedar Hill (K75), Clear Crk Cyn (KU), Hardup (K75), Kelton Pass (K70), Man- tua, Portage (KU), Snowville (K70), Wellsville Mts (KU), Wildcat Hills (K75), Willard Basin (F). CACHE: Ant Valley, Antelope Valley, Bear River Mts, Beaver Crk, Blacksmith Fk Cyn, Elk Valley, Franklin Basin, Green Cyn, Hodges Cyn, Leeds Cyn, Logan (KU), Logan Cyn (C42), Mendon Cold Spngs, River Heights, Millville, Rock Crk, Tony Grove (KU). EMERY: Ferron (RG), Hideout Cyn nr Green River (U). GARFIELD: Boulder (U), Bryce Cyn Nat Park (WU), Carcass Crk (on Boulder Mt) (U). IRON: Cedar Breaks Nat Mon (F), Cedar City 19 mi E (A). JUAB: Indian Farm Cyn (U). KANE: Cedar City 24.3 mi E (A), Glendale (159), Kanab Cyn (C42). RICH: Garden City, Meadowville and 8 mi NW, Monte Cristo (KU). SALT LAKE: Big Cottonwood Cyn, Brigh- ton, Butterfield Cyn, Mt Olympus, Red Butte Cyn (U), Salt Lake City (C42). SAN JUAN: Abajo Mts (G58), La Sal Crk (RG), Monticello (U). SANPETE: Bluebell Flats (KU), Ephraim Cyn (U), Pleasant Crk (BAD). SUMMIT: Kamas 21 mi E, Mirror Lake 1 and 6.4 mi N (A), Soap- stone Ranger Sta (U), Wyoming brdr on U150 (A). TOOELE: Clover, Fisher Pass, Grantsville, Tooele (C42), S Willow Cyn (U). UINTAH: Ashley Crk nr Ver- nal (U), Dry Fk rd 15 and 22.8 mi N U121 (A). UTAH: American Fk Cyn (U), Aspen Grove (BY), Halls Fk rd 5.2 mi N from Hobble Crk rd (A), Provo (BY), Silver Lake Flat (A). WASATCH: Francis 8.1 mi E (A), Heber (U). WASHINGTON: Kolob (159), Pine Valley (City) (BAD). WEBER: Beaver Crk, Lime Spngs, Monte Cristo 6 mi S (KU), Ogden (C42). COUNTY UNKNOWN: Current Crk (C42) (? = Currant Crk either in Duchesne, Utah, or Wasatch Co). Smith (1979:1452) lists this as an eastern to western species including Arizona, which nests in a variety of situations in soil and un- der rocks and logs in forests or open areas. Gregg (1963:517) lists it from Colorado be- tween 5154 and 12,500 ft under rocks and logs and in thatched domes in a variety of habitats, predominantly in conifers, oak, and pinyon-juniper. Wheeler and Wheeler (1978:395) found it between 6100 and 11,500 ft in Nevada, and nesting frequently in wood in North Dakota (1977). Allred and Cole December 1982 Allred: Ants of Utah 465 (1971:239) found it in Idaho in an association of juniper-rabbitbrush-winterfat-sagebrush- grass. Grundmann (1958:165) designated it as a stream-side species in Utah between 4500 and 10,000 ft under stones or in low irregular moimds with numerous openings. Ingham (1959) found it imder stones and logs in as- pen, fir, spruce, and bristlecone pine in southern Utah. Cole (1966:23) found it in southern Nevada mider stones restricted to pinyon-juniper, under stones and logs, and sometimes in craters or small earthen mounds at rather high elevations in Utah (1942:381). Knowlton (1975:2) found it associated with grass, sagebrush, shadscale, and rabbitbnish in northern Utah. There were 122 ants in eight collections taken from under rocks. In one collection it was under the same rock with Formica ar- gentea, once F. gnava, once F. podzoHca, once F. podzolica and Solenopsis molesta, and once F. gnava, Mymiica enienjana, and M. monticola. In the association with gnava and Mymiica, when the rock was removed and the ants disturbed, some began pulling on the legs of others. In the association with pod- zolica and Solenopsis, the earth under the huge rock was divided into separate con- spicuous dwelling levels in which the For- mica occurred. The smaller Solenopsis were in a separate burrow. Thirty-eight ants in one collection were in a burrow with Pohjgerns breviceps under a log. Forty-one ants in three collections were found vmder logs, once mi- der the same log with Formica gnava and F. podzolica, and once with F. siihnuda and Lasius alienus. One ant was found in the open crawling on the ground. Six collections were in aspen: two in association with grass, sagebrush, and pine; one grass, herbs, and pine; and one pine. Three collections were in conifers; two in a grass, sedge, and herb meadow; one sagebrush; and one cotton- woods. In 82 recorded Utah habitats 49 were in montane areas. In 45 known elevations be- tween 4288 and 10,500 ft, 20 were under 6000 and 3 were over 10,000. Beck et al. (1967:69) found it feeding on dead rodents in two instances in Utah. Formica gnava Buckley F. gnava Buckley, 1866, Proc. Ent. Soc. Phila. 6:156; Smith 1979:1453. F. forcliana: Rees and Grundmann 1940:9; Cole 1942:380. F. rufibarhis gnava: Rees and Grundmann 1940:10; Cole 1942:383; Creighton 1950:539; Ingham 1959:82; Allred and Cole 1979:98. Records (Map 8): DUCHESNE: Avintaquin Cmpgnd (A). GARFIELD: Boulder Mt (U). GRAND: Dewey (U). IRON: Cedar City 19 mi E (A). KANE: Glen Cyn City (AC). MORGAN:'Morgan (C42). PIUTE: Marysvale 4.1 mi S (A). RICH: Woodruff 18.4 mi W (A). SALT LAKE: Big Cottonwood Cyn (U). SAN JUAN: Rainbow Bridge Nat Mon (F). SANPETE: Orangeville 23.9 mi W, Sky- line Drive 1 mi N Jet Orangeville and Ephraim rds (A). SUMMIT: Kamas 26.4 and 28.5 mi E, Mirror Lake 1 and 17.3 mi N (A), Park City (F). UINTAH: Red Cloud Loop rd 14 mi W U44 (A). UTAH: Halls Fk rd 5.2 mi N Hobble Crk rd (A), Lehi (RG), Nebo Loop rd 0.8 mi S Santaquin Cyn, Payson Cyn 12.3 mi up, Santaquin 6.7 mi E (A). WASATCH: Hanna 14.3 mi W (A). WASH- INGTON: Snow Cyn (159). WEBER: Ogden Cyn (C42). Smith (1979:1453) lists this species from western United States, including Utah, Colo- rado, Arizona, and Nevada, nesting under rocks in desert areas or open woods. Gregg (1963:539) lists it between 5333 and 9000 ft under rocks and in thatched nests pre- dominantly in pinyon-juniper-oak habitats in Colorado. Cole (1942) indicates its habitat in Utah as under stones or nests without craters in open areas. Ingham (1959, 1963) found it in soil-lacking mounds in sand sage in south- ern Utah. Allred and Cole (1979:98) found it in juniper-ephedra-grass and ephedra-grass habitats in southern Utah. There were 281 ants in 11 collections taken from under logs. In one collection it was under the same log as Formica fusca and F. podzolica, once with F. subnuda, once F. neoclara and Camponotus modoc, once C modoc, and once Myrmica brevispinosa. Ninety ants in 2 collections were taken from inside a log. There were 131 ants in 7 collec- tions taken from under rocks. In one collec- tion gnava was under the same rock as F. fusca; once with fusca, Myrmica emery ana, and M. monticola; once F. podzolica; and once F. neogagates, F. obtusopilosa, and F. perpilosa. When the rock covering gnava and fusca was removed, some ants were seen pulling others by their legs. The ants being pulled seemed not to demonstrate a defensive or aggressive behavior to the "pullers," sug- gesting a captive arrangement rather than fighting between different colonies. Ten ants in one collection were found under a slab of bark on the ground, and one ant was taken in 466 Great Basin Naturalist Vol. 42, No. 4 the open. Immatures were found under one rock. Eleven collections were in aspen: 4 in association with fir, 3 with other conifers, one sagebrush and conifers, and one grass, herbs, and fir. Six collections were in co- nifers, 4 grass-herb meadows, and one sage- brush. In 26 recorded Utah habitats it was taken 18 times in montane forest. In 10 known elevations between 2900 and 10,500 it was taken 6 times under 5000, once over 7000 ft. Formica haemorrhoidalis Emery F. nifa integra var. haemorrhoidalis Emery, 1893, Arb. Zool. Jahrb. Syst. 7:644. F. truncicola integroides var. haemorrhoidalis: Rees and Grundmann 1940:8. F. riifa liaemorrhoidalis: Cole 1942:381. F. integra liaemorrhoidalis: Creighton 1950:488; Beck et al. 1967:70. F. haemorrhoidalis: Knowlton 1970:209, 1975:3. Records (Map 9): BOX ELDER: Hansel (KU), Kelton (K70) and 9 mi N (KU), Snowville (K75), Trematon (KU) (? = Tremonton). CACHE: Avon, Bear River Mts, Green Cyn, Elk Valley, Leeds Cyn (KU), Hyde Park (C42), Logan Cyn, Smithfield Cyn (KU). CARBON: My- ton rd 15 mi E US6 (A), Scofield (BAD). DAGGETT: Deep Crk (BAD). DAVIS: KaysviUe (C42). DUCH- ESNE: Duchesne 12.4 mi S (A). GARFIELD: Bryce.Cyn Nat Park (RG). JUAB: Callao 5 mi E(A). RICH: Chalk Crk (U). SAN JUAN: Blanding, Monticello (U). SUM- MIT: Kamas 9.2 mi E (A). TOOELE: Mercur (BAD). UTAH: Eureka 0.5 mi E (A), Spanish Fk Cyn (KU). WASATCH: Midway 3.7 mi NW (A). Smith (1979:1458) lists this species from midwest to western United States, including Colorado, nesting under logs or stones in areas of moderate to sparse cover. Gregg (1963:556) lists it between 5100 and 10,000 ft under rocks and logs in thatched nests pre- dominantly in conifer habitats in Colorado. Allred and Cole (1971:239) found it in Idaho abundantly in an association of wild rye- grass, moderately so in rabbitbrush-sage- brush-grass, and rarely in rabbitbrush-sage- brush-grass-winterfat. La Rivers (1968:9) lists it from Nevada. Wheeler and Wheeler (1977) found it frequently in thatched mounds in North Dakota. Nests in Utah are under logs or stones, usually with a scattering of detritus on the periphery (Cole 1942:381). Knowlton (1975:3) found it associated with sagebrush in northern Utah. Thirty-two ants in three collections were found under rocks. Once it was under the same rock as Formica integroides, and once with Solenopsis molesta. In the latter case the burrows of the two species were separate. Forty ants were taken from a mound of soil covered with a layer of sticks. Three ants in two collections were taken in the open. When disturbed in their colony these ants frequently rear back on their 2nd and 3rd legs, tucking the abdomen underneath, leav- ing the front legs and mandibles in an up- ward, apparently defensive position. Four collections were in sagebrush: one in associ- ation with herbs; one grass and herbs; one grass, herbs, serviceberry, cliffrose, juniper, and pinyon; and one aspen and conifers. One collection was in greasewood, and one in oak. In 28 recorded Utah habitats it was taken 13 times in montane forest. In eight known ele- vations between 4225 and 7977 ft it was found most frequently below 5000 and above 6000. Beck et al. (1967:70) found it feeding on dead rodents in four instances in Utah. Formica hewitti Wheeler F. hewitti Wheeler, 1917, Proc. Amer. Acad. Arts Sci. 52:552; Creighton 1950:533; Smith 1979:1453. Records: BOX ELDER: Locomotive Spngs (KU). CACHE: W Hodges Cyn (KU). GRAND: Warner Ranger Sta (F). Smith (1979:1453) lists this species from eastern to western United States, including Utah, nesting under rocks or wood in woods and forests. Gregg (1963:526) hsts it from Colorado between 5354 and 10,000 ft under rocks and logs predominantly in conifer habi- tats. Wheeler and Wheeler (1978:395) found it between 6700 and 11,600 ft in Nevada. All- red and Cole (1971:239) found it in Idaho in an association of rabbitbrush-sagebrush-grass- winterfat. One collection in Utah was at 9750 ft. Formica integroides Emery F. rufa obscuriventris var. integroides Emery, 1893, Zool. Jahrb. Syst. 7:644. F. truncicola integroides: Rees and Gnmdmann 1940:8. F. trunciola integroides var. coloradensis: Rees and Gnmdmann 1940:8. F. integroides: Rees and Grundmann 1940:8. F. rufa coloradensis: Cole 1942:381. F. integroides coloradensis: Creighton 1950:489; Ingham 1959:79; Smith 1979:1458. F. integroides planipilis: Knowlton 1970:209, 1975:5. F. integroides propinqua: Smith 1979:1459. Records (Map 9): BOX ELDER: Hansel Mts, Locomo- tive Spngs, Snowville, Wellsville Mts, Willard Basin December 1982 Allred: Ants of Utah 467 (KU). CACHE: Bear River Mts, Blacksmith Fk Cyn, Franklin Basin, Leeds Cyn (KU), Logan Cyn (RG), Men- don Cold Spng, Monte Cristo (KU), Providence (C42), Tony Grove Cyn, W Hodges Cvn (KU). CARBON: My- ton rd 15 mi E US6 (A). GRAND: Warner Ranger Sta (Gr63). JUAB: Eureka 0.5 mi E (A), McClellan Lake, NE Nephi (KU). KANE: Duck Crk Ranger Sta (C42). MIL- LARD: Swasev Spngs (RG). RICH: Randolph, Sage Crk (KU), Utah-Wyoming brdr 0.5 mi S on U16 (A). SALT LAKE: Big Cottonwood Cyn (U). SANPETE: Wales 3.3 mi W (A). UINTAH: Dry Fk rd 22.8 mi N U121 (A), Paradise Park 11 mi S (U)'. UTAH: Thistle 14.6 and 20.4 mi E, Tibbie Fk Cyn (A). WASATCH: Daniels Cyn (RG). WASHINGTON: Kolob (159). WEBER: Hooper, Slaterville (C42). Creighton (1950) lists six and Smith (1979:1458) five subspecies from western United States, including Utah, Colorado, Ne- vada, and Idaho, nesting under logs that are usually thatched. The four that have been re- corded from Utah may be separated by the following key. 1. Erect hairs other than double row on middle and hind tibiae abundant 2 — Erect hairs consist only of a double row on flexor surface 3 2(1). Head and thorax clear red, legs about same color as thorax coloradensis — Head and thorax of smaller workers marked with brown, legs brownish black ... planipilis 3(1). Occipital angles of head with erect hairs integroides — Occipital angles of head lack erect hairs propinqua Gregg (1963) lists this species between 5354 and 12,000 ft under rocks, logs, and in thatched nests predominantly in conifer habi- tats in Colorado, and lists a record for Utah. Cole (1966:23) found it in southern Nevada only in pinyon-juniper, nesting in thatched mounds adjacent to shrubs. It also nests under logs with an adjacent thatching of detritus, resulting in dome-shaped mounds in Utah (Cole 1942:381). Wheeler and Wheeler (1978:394) found it between 5600 and 10,200 ft in Nevada. Ingham (1959) found it in mounds of detritus in sagebrush and oak in southern Utah. Knowlton (1975:5) found it in thatched nests in northern Utah. Fifty ants in four collections were taken from mounds of sticks, one of them mixed with soil next to a log. In this latter case, ants of Formica obscuripes and F. obscuriventris were also present. In two other mounds ants of F. oreas were present. Ninety ants in three collections were taken from under rocks, once under the same rock with F. laeviceps and once with F. haemorrhoidalis. Four ants in one collection were found crawling in the open. Immature stages were found under two rocks in late June and early July. Six collec- tions were in sagebrush: one in association with grass; one grass and herbs; one grass, herbs, and rabbitbrush; one grass, clover, and Russian thistle; one grass, shrubs, juniper, and pinyon; and one herbs, rabbitbrush, maple, and oak. One collection was in aspen. In 35 recorded Utah habitats it was taken 19 times in montane areas. In 14 known elevations be- tween 4240 and 8555 ft it was collected most frequently under 5000 and over 8000. Formica laeviceps Creighton F. rufa laeviceps Creighton, 1940, Amer. Mus. Nat. Hist. Nov. 1055:7; Cole 1942:380. F. laeviceps: Creighton 1950:491; Smith 1979:1459. Records (Map 9): BOX ELDER: Snowville (US). CACHE: Hyde Park (KU). DAVIS: Kaysville (US). GRAND: Warner Ranger Sta (C42). JUAB: Nebo Loop rd 9.9 mi S Santaquin Cyn (A). SALT LAKE: Big Cot- tonwood Cyn, Mt Dell Res (U). SANPETE: Wales 3.3 mi W (A). SUMMIT: Chalk Crk (U). TOOELE: Tooele Cyn (U). WASHINGTON: Pine Valley (City) (US). WE- BER: Slaterville (US). Smith (1979:1459) lists this species from midwest and western United States, including Utah and Colorado, nesting under stones and logs with little debris in open areas. Gregg (1963:564) lists it between 5200 and 8500 ft in sagebrush and grass habitats in Colorado. Cole (1942:380) indicates that it nests under stones and logs in areas of moderate to sparse cover in Utah. Ten ants in one collection were taken from under a rock in association with Formica in- tegroides. Twenty ants in two collections were taken crawling on the ground in open 468 Great Basin Naturalist Vol. 42, No. 4 areas. Three collections were in sagebrush, two in association with rabbitbrush. In 12 re- corded Utah habitats it was taken 7 times in montane forest. In 8 recorded elevations be- tween 4240 and 9750 ft it was taken most frequently under 5000, once over 8000. Formica lasioides Emery F. lasioides Emery, 1893, Zool. Jahrb. Syst. 7:646. F. neogagates lasioides var. vetttla: Rees and Grundmann 1940:10; Cole 1942:384. Records (Map 9): BOX ELDER: Wellsville Mts (KU). CACHE: Ant Valley, Antelope Valley, Beaver Mt, Blacksmith Fk Cyn, Elk Basin, Franklin Basin, Green Cyn, W Hodges Cyn, Hymm, Leeds Cyn, Logan Cyn, Millville, Monte Cristo, Pole Crk Spng, Ricks Spng, Tony Grove (KU). EMERY: Ferron Res, Green Rive'r, Gunnison Butte (RG). GRAND: Warner Ranger Sta (C42). JUAB: Deep Crk Mts (U). RICH: Garden City (KU), Randolph, and 8 and 10 mi S, Sage Crk (KU). SALT LAKE: Big Cottonwood Cyn (U). SANPETE: Ephraim Cvn, M^ajors Flats (KU). SUMMIT: Mirror Lake 11.6 mi N (A). UINTAH: Bonanza, Elk Basin, Hy- mm, Millville (KU). UTAH: Nibley (KU), Payson Cyn 7.3 mi up (A). WASATCH: Strawberry Res 4 mi S (A). WASHINGTON: Pine Valley Cmpgnd (KU). WEBER: Beaver Crk (KU), Wolf Crk (U). COUNTY UNKNOWN: Beaver Head (KU). Smith (1979:1449) lists this species from midwest to western United States, including Colorado and Arizona, nesting under stones or in soil or small craters. Gregg (1963:500) lists :^ between 4800 and 10,505 ft under rock.-, and wood in a variety of habitats in Colorado. Cole (1966:23) found its nests in southern Nevada under stones in pinyon-juni- per, and indicated that some ants construct small crat.xs in Utah (1942:384). Wheeler and Wheeler (1978:393) found it between 6200 and 10,900 ft in Nevada, and frequently under rocks in North Dakota (1963). Allred and Cole (1971:239) found it in Idaho in asso- ciations of wild rye-grass and rabbitbrush- sagebrush. Twenty ants in two collections were taken from under rocks, once under the same rock with Formica argentea and once with F. oh- scuriventris. Four ants in one collection were taken singly in the open. Three collections were in sagebrush: one in association with le- gumes and two with aspen and conifers. In 39 recorded Utah habitats it was taken 26 times in montane forest. In 13 known eleva- tions between 4087 and 11,000 ft it was taken most frequently under 6000, once over 10,000. Formica litnata Wheeler F. limata Wheeler, 1913, Bull. Mus. Comp. Zool. 53:400; Creighton 1950:458; Beck et al. 1967:70; Smith 1979:1450. Records (Map 10): BOX ELDER; Raft River Mts (U). KANE: Adairville (BAD). SAN JUAN: Pack Crk (KU). SEVIER: Fremont 9.7 mi N (A), Koosharem (BAD), Richfield 2.3 mi S (A). TOOELE: Dugway 11.9 mi E (A). WASATCH: Francis 8.1 mi E (A). WEBER: Woodruff 34.8 mi W (A). Smith (1979:1450) lists this species from midwest and western United States, including Utah, Colorado, and Nevada, nesting under stones or in craters in grasslands. Gregg (1963:502) lists it between 5000 and 9700 ft under rocks and logs in a variety of habitats in Colorado. Wheeler and Wheeler (1963) found it frequently under rocks in North Da- kota. Cole (1966:24) found its nests under stones in pinyon-juniper in southern Nevada. Sixty ants in three collections were taken from under rocks. In one case they were un- der the .same rock with Formica neoclara, once F. occulta, and once F. podzolica. Nine ants in two collections were taken in the open on the ground. Immatures were found under one rock in early August. Two collec- tions were in sagebrush, one associated with matchbmsh. One collection was in Russian thistle, one cottonwoods, and one juniper and pinyon. In nine recorded Utah habitats it was found four times in montane forest. In three collections it was taken at 5350, 5375, and 6850 ft. Beck et al. (1967:70) found it feeding on dead rodents in two instances in Utah. Formica manni Wheeler F. manni Wheeler, 1913, Bull. Mus. Comp. Zool. 53:389; Rees and Grundmann 1940:8; Cole 1942:378; Ing- ham 1959:77; Knowlton 1970:209; 1975:3; Smith 1979:1450. Records (Map 10): BOX ELDER: Cedar Hill (US), Curlew Jet (K70), Hansel Mts (K75), Hardup (C42), Kel- ton (K75), 5 mi N (US) and 9 mi NW (KU), Kelton Pass (K70), Lampe (C42) (? = Lampo), Locomotive Spngs (K75), Promontory (KU), Snowville (K70), 6 mi W (KU) and 17 mi SW (KU), Wildcat Hills (US), Willard (C42). CACHE: Green Cyn (KU), Logan (C42). MILLARD: Tule Spngs (C42). RICH: Allen Cyn (KU). SALT LAKE: Wasatch Mts (RG). TOOELE: Stansbury Island (C42). UTAH: Jordan Narrows (C42), Silver Lake Flat (A). WASATCH: Kamas 11 mi SE (U). WASHINGTON: Leeds (RG), St George (KU). Smith (1979:1450) lists this species from western United States, including Utah, Ne- vada, and Idaho, nesting under stones in December 1982 Allred: Ants of Utah 469 desert areas. Allred and Cole (1971:239) foimd it in Idaho in an association of sage- brush-grass, rarely in other shrub types. Cole (1942:379) indicates its habitat as under stones in Utah. Ingham (1963) found it in jimiper, sagebrush, galletagrass, rabbitbnish, winterfat, shadscale, and greasewood in southern Utah. Knowlton (1975:3) found it associated with rabbitbrush in northern Utah. One specimen was taken from under a log in grass of an open area of aspen. Twenty- one ants of Formica suhniida were under the same log. In 27 recorded Utah habitats it was taken five times in montane areas. In 13 known elevations between 2750 and 9000 ft it was taken most frequently between 4000 and 5000, twice over 5000. Formica microgyria Wheeler F. microgijna Wheeler, 1903, Bull. Amer. Mus. Nat. Hist. 19:645; Creighton 1950:504; Gregg 1963:586; Knowlton 1975:3; Smith 1979:1462. Records: BOX ELDER: Hansel Mts (KU), Snowville (K75). CACHE: Leeds Cyn (KU). GRAND: Warner Ranger Sta (Gr63). Smith (1979:1462) lists this species from western United States, including Utah, Colo- rado, Nevada, and Wyoming, nesting under thatched stones in meadows and open forest. It is frequently associated with Formica ar- gentea, F. lasioides, and F. neogagates. Gregg (1963:586) lists it between 5800 and 9400 ft under rocks and logs and in thatched nests predominantly in conifer habitats in Colo- rado, and gives a record for Utah. Cole (1966:24) found one worker in pinyon-juniper in southern Nevada. One collection in Utah was taken at 4544 ft. Formica mucescens Wheeler F. truncicola mucescens Wheeler, 1913, Bull. Mus. Comp. Zool. 53:392; Creighton 1950:491. F. riifa mucescens: Cole 1942:380. F. mucescens: Smith 1979:1460. Records: GARFIELD: Bryce Cvn Nat Park (C42). SANPETE: Wales 3.3 mi W (A). Smith (1979:1460) lists this species from Utah and Colorado nesting under stones in open areas. Gregg (1963:566) lists it between 5500 and 8750 ft imder rocks and in thatched nests in grass in conifer habitats in Colorado. Cole (1942:380) indicates its habitats in Utah as under stones and logs. Four ants in one collection were taken un- der a rock in an association of sagebrush, rab- bitbrush, maple, and oak. In one collection it was taken at 7977 ft. Formica neoclara Emery F. fusca var. neoclara Emery, 1893, Arb. Zool. Jahrb. Syst. 7:646; Cole 1942:383. F. neoclara: Creighton 1950:535; Beck et al. 1967:70; Knowlton 1970:209, 1975:3; Francoeur 1973:92. F. pruinosa: Beck et al. 1967:70; Knowlton 1970:210, 1975:4. Records (Map 10): BEAVER: Beaver (U), Beaver Cyn (F). BOX ELDER: Bear River City, Brigham, Corinne, Garland (US), Hansel Mts (US), Hardup, Kelton (K70) and 6 mi N (KU), Lucin (BAD), Mantua (F), Morton Cold Spngs (KU), Newton (F), Riverside (US), Snowville (K70), Tremonton (F), Willard (US), Willard Basin (KU). CACHE: Amalga (US), Ant Valley, Blacksmith Fk Cyn, Carter Crk, Clarkston, Franklin Basin, W Hodges Cyn (KU), Hyde Park (F), Lewiston, Logan (C42), Logan Cyn (KU), Mendon, Millville, Monte Cristo, Newton (KU), Paradise (US), Providence (KU), Smithfield (C42), Smithfield Cyn, Tony Grove (KU), Trenton (F), Wells- ville (C42). CARBON: Clear Crk Cyn, Price (F) and 2 mi S (U). DAVIS: Farmington (F), Kaysville (US), Lay- ton (F). DUCHESNE: Lake Atwood, Roberts Pass (BY). EMERY: Hideout Cyn nr Green River (U), Huntington (KU). IRON: Paragonah (U). JUAB: Topaz Mt (KU). KANE: Alton, OrderviUe (F). MORGAN: Morgan (BAD), Porterville (KU). RICH: Allen Cyn (KU), Chalk Crk (U), Laketown, Monte Cristo (KU).' SALT LAKE: Brighton (C42), Hunter, Murrav (US), Salt Lake City (C42). SANPETE: OrangeviUe 23.9 mi W (A), Palisade (KU). SEVIER: Aurora (F), Richfield 2.3 mi S (A). SUM- MIT: Kamas 11 mi E (U), Park Citv (F), Wanship (US). TOOELE: Grantsville (US). UINTAH: Bonanza (KU). UTAH: Lehi (F), Provo (A), Thistle (F), and 7.7 mi E (A), Utah Lake (F). WASATCH: Bald Mt (KU), Francis 8.1 mi E, Hanna 3.6 mi W (A), Heber (F). WASHING- TON: locality unknown (F). WAYNE: Hanksville (BAD). WEBER: Ogden (F), Riverdale (US), Slaterville (F). Smith (1979:1454) lists this species as east- ren and western United States (no inter- mountain state is listed), where it nests in the soil, sometimes with loose mounds of soil and detritus, grasslands, and open woods. Hunt and Snelling (1975:23) list it from Arizona. Gregg (1963:529) lists it from Colorado be- tween 3500 and 9000 ft under rocks and logs in a variety of habitats, predominantly in cot- tonwood-willow areas. La Rivers (1968:10) lists it from Nevada. Wheeler and Wheeler (1977) found it frequently in earthen mounds, also under rocks and wood in North Dakota. Cole (1942:383) indicates its habitat in Utah as soil, with numerous entrances in crude, flat, confluent craters. Knowlton (1975:3) found it associated with sagebrush and rab- bitbrush in northern Utah. 470 Great Basin Naturalist Vol. 42, No. 4 Eighty ants in four collections were taken from under rocks. In one case they were un- der the same rock with Formica pallidefulva and once with F. limata. Ten ants in one col- lection were taken under the same log with F. gnava and Camponotus modoc. Fifteen ants in three collections were taken singly in open areas. Eggs were found under one rock in early July, and in mid-July many winged forms were found under a rock. These tried to hide and escape when disturbed. Two col- lections were in cottonwoods; one grass, herbs, and sagebrush; one herbs adjacent to a cultivated area; one Russian thistle; and one aspen and conifers. In 87 recorded Utah habi- tats it was taken 26 times in montane areas. In 49 recorded elevations between 4125 and 11,000 ft it was taken most frequently (44 times) under 6000. Beck et al. (1967:70) foimd it feeding on dead rodents in one in- stance in Utah. Formica neogagates Emery F. fusca subpolita var. neogagates Emery, 1893, Zool. Jahrb. Syst. 7:646. F. neogagates: Rees and Grundmann 1940:10; Cole 1942:384; Hayward 1945:120; Creighton 1950:459; Ingham 1959:76; Knowlton 1970:209, 1975:3. Records (Map 11): BOX ELDER: Beaver Dam (KU), Cedar Crk (City) (K70), Cedar Hill (KU), Curlew Jet (US), Hansel Mts (K75), Hardup (US), Kelton (K75), Kel- ton Pass, Locomotive Spngs (K70), Promontory Pt (C42), Snowville (K70) and 17 mi SW (KU), Taylor Farms (K75), Wildcat Hills (US). CACHE: Avon, 'Blacksmith Fk Cyn, Clarkston, Elk Valley, Green Cyn (KU), Logan, Logan Cyn (C42), Monte Cristo, Paradise, Petersboro, Providence, Tony Grove (KU). CARBON: Helper (KU). EMERY: Hideout Cyn nr Green River (U). GARFIELD: Boulder (U), Jet U12 and U63 0.5 mi E(A). IRON: Ka- naraviUe (US). JUAB: Callao 5 mi E (A), Mt Nebo, To- paz Mt (KU). MILLARD: Tule Spngs, White Valley (C42). PIUTE: Marysvale 4.1 mi S (A). RICH: Monte Cristo, Randolph 10 mi SW (KU), Sage Crk Jet 5.1 mi W, Woodruff 4 mi W, Wyoming brdr 0.5 mi S on U16 (A). SALT LAKE: Alta, Cottonwood Cyn (C42), Red Butte Cyn (U), Salt Lake City (C42). SANPETE: Or- angeville 19.5 mi W (A). SEVIER: Salina, Salina Cyn nr Fremont Jet (U), Sevier (C42). TOOELE: Lake Point (C42). UTAH: Aspen Grove (BY), Nibley, Spanish Fk Cyn (KU). WASATCH: Provo River N Fk (U). WASH- INGTON: Hurricane (US), Kolob (159), La Verkin (US), Rockville (KU), Santa Clara (US), St George (KU), Virgin River E Fk (159). WEBER: Beaver Crk, Monte Cristo 6 mi S, Thomas Spng (KU). Smith (1979:1450) lists this species from midwest to western United States, including Arizona, nesting under stones or in the open with or without a mound or crater in grass- lands and dry situations. Gregg (1963:505) lists it from Colorado between 3500 and 9700 ft under rocks and in soil hummocks in a va- riety of habitats. AUred and Cole (1971:239) found it in Idaho in associations of wild rye- grass and rabbitbrush-sagebrush-grass-winter- fat. In southern Nevada Cole (1966:26) found its nests under rocks in pinyon-juniper, and in Utah in small craters (1942:384). Wheeler and Wheeler (1963) found it frequently under rocks in North Dakota. Ingham (1959, 1963) found it under rocks and in small craters in open areas in oak, sagebrush, cultivated areas, shadscale, and greasewood in southern Utah. Knowlton (1975:3) found it associated with sagebrush, rabbitbrush, and sunflowers in northern Utah. Seventy-eight ants in three collections were taken from moundless burrows. Twenty-one ants in two collections were taken from under rocks, once under the same rock with Formica gnava, F. ohscuripes, and F. perpilosa. Nineteen ants in two collections were taken in the open. Five collections were in sagebrush: one in association with grass, one grass and herbs, and one grass and rabbit- brush. One collection was in grass and one in greasewood. In 65 recorded Utah habitats it was found 19 times in montane forest. In 32 recorded elevations between 2625 and 8585 ft it was taken most frequently between 4000 and 5000. Formica neorufibarbis Emery F. fiisca var. neorufibarbis Emery, 1893, Zool. Jahrb. Syst. 7:646; Rees and Grundmann 1940:9; Cole 1942:382. F. fusca gelida: Rees and Grundmann 1940:9; Cole 1942:382; Hayward 1945:120. F. neorufibarbis: Grundmann 1958:166; Knowlton 1970:209, 1975:3; Francoeur 1973:225, 244. F. neorufibarbis gelida: Beck et al. 1967:70. Records (Map 11): BOX ELDER: Bear River City, Brigham, Collinston, Corinne, Curlew, Fielding, Gar- land, Hardup, Park Valley (City), Promontory Ridge, Riverside ( = Belmont), Snowville, Thatcher, Tremon- ton, Willard (C42), Willard Basin (KU). CACHE: Amalga (C42), Franklin Basin (KU), Hyde Park, Lewis- ton, Logan, Logan Cyn (C42), Mendon, Monte Cristo (KU), Newton, Paradise, Petersboro, Richmond (C42), Ricks Spng (KU), Sardine Cyn, Smithfield (C42), Tony Grove Lake (KU), Trenton (C42), Wellsville (RG). DAG- GETT: Green Lake (RG). DAVIS: Farmington, Farm- ington Cyn (KU), Kaysville (C42), Woods Cross (RG). DUCHESNE: Fniitland 4 mi W (U), Mirror Lake (KU). December 1982 Allred: Ants of Utah 471 GARFIELD: Osiris (U). GRAND: Moab (G58). IRON: Cedar City, Parowan, Summit (C42). JUAB: Callao (BAD). KANE: OrderviUe (G58). MILLARD: Holden, Leamington (C42). RICH: Allen Cyn (KU), Laketown (C42), Monte Cristo (KU). SALT LAKE: Butterfield Cvn, Hmiter, Little Cottonwood Cyn, Mill Crk Cyn, Mt Dell Res (U), Murray (C42), Parleys Cyn (RG), Red Butte Cyn (U), Salt Lake City (C42). SAN JUAN: Angels Pk (KU)! Blanding (G58), Geyser Pass (BAD), Monticello (G58). SANPETE: Ephraim (RG), Ephraim Cyn (KU). SEVIER: Salina (C42). SUMMIT: Henrys Fk Basin (RG), Kamas (C42), Park City (F), Wanship (C42). TOOELE: Fisher Pass, Grantsville, losepa, Orrs Ranch, St Johns (C42). UINTAH: Gusher (C42), Ouray (C42), Paradise Park, Vernal (U). UTAH: American Fk Cyn (U), Emer- ald Lake (BAD), Hobble Crk, Lehi, Orem, Provo (C42). WASATCH: Daniels Cyn, Deer Crk Res, Heber (U), Lost Lake (BY), Midway (U), Soapstone Cyn (RG). WE- BER: Hooper, Ogden, Riverdale, Roy (C42). COUNTY UNKNOWN: Sharon (C42). Smith (1979:1454) lists this species from midwest to western United States, including Arizona, nesting in rotting wood or under rocks in montane forest. Gregg (1963) lists it from Colorado between 6900 and 11,542 ft under rocks and logs and in hummocks pre- dominantly in conifer habitats. Cole (1966:26) foimd its nests in southern Nevada under stones in pinyon-juniper, also in logs in wooded areas in Utah (1942:382). Wheeler and Wheeler (1978:395) found it between 6200 and 12,200 ft in Nevada, frequently in earthen mounds and under rocks and wood in North Dakota (1963). Grundmann (1958:166) indicates it as an inhabitant of cultivated and stream side areas usually below 5000 ft in Utah. In 100 recorded Utah habitats 27 were in montane forest. In 71 elevations between 4042 and 11,300 ft, 52 were under 6000, 14 between 6000 and 8000, and one over 11,000. Beck et al. (1967:70) found it feeding on dead rodents in three instances in Utah. Formica obscuripes Forel F. nifa obscuripes Forel, 1886, Ann. Ent. Soc. Belg. 30:39; Rees and Grundmann 1940:8; Cole 1942:380. F. rufa aggerans: Rees and Grundmann 1940:8. F. ritfa melanotica: Cole 1942:380. F. obscuripes: Ingham 1959:80; Gregg 1963:570; Beck et al. 1967:70; Knowlton 1970:210, 1975:3; Smith 1979:1460. Records (Map 12): BOX ELDER: Bear River City, Bear River Cyn, Brigham, Collinston, Garland, Honey- ville (C42), Hansel Mts, Hardup, Kelton, Kelton Pass (K70), Promontory Pt, Raft River Mts (U), Snowville (K70), WellsviUe Mts, Wildcat Hills (KU). CACHE: Amalga (C42), Ant Valley, Blacksmith Fk Cyn, Franklin Basin, W Hodges Cyn (KU), Logan, Logan Cyn (C42), Monte Cristo (KU), Providence, Smithfield (C42), Tony Grove Lake (KU). DAGGETT: Green Cyn (RG), Rado- sovich Ranch (BAD). DAVIS: Farmington, Layton (C42). EMERY: Carter Crk nr Green River (U). GRAND: La Sal Mts (BY), Warner Ranger Sta (Gr63). JUAB: McClellan Lake (KU), Nebo Loop rd 9.9 mi S Santaquin Cyn (A), Trout Crk (City) (C42). MILLARD: Swasey Spngs (RG). MORGAN: Morgan (C42). PIUTE: Marysvale 4.1 mi S (A). RICH: Meadowville 8 mi NW (KU), Monte Cristo Ranger Sta, Randolph (BAD) and 10 mi SW, Sage Crk (KU). SALT LAKE: Big Cottonwood Cyn (U), Holladay, Midvale, Murray, Parleys Cyn (C42), Red Butte Cyn (U). SANPETE: Fountain Green (C42), Wales (RG).' SEVIER: Monroe Mt (BAD). TOOELE: Stansbury Island (C42), S Willow Cyn (U). UINTAH: Paradise Park 11 mi S (U). UTAH: Aspen Grove (BY), Lehi (RG), Provo (C42), Spanish Fk (BY), Spanish Fk Cyn (KU), Tibbie Fk Cyn (A). WASATCH: Kamas 8 mi S (U). WASHINGTON: Harrisburg (C42), Pine Valley (City) (KU). WAYNE: Capitol Reef Nat Park (U). WE- BER: Harrisville, Hooper, Huntsville, Ogden, Slaterville (C42), Woodruff 34.8 mi W (A). Smith (1979:1460) lists this species from midwest to western United States, including Utah, nesting in large mounds of detritus in open areas. It is a known predator of the pine sawfly. Gregg (1963:570) lists it from Colo- rado between 3500 and 9500 ft under rocks and logs and in thatched domes in a variety of habitats, and gives a record for Utah. La Rivers (1968:10) lists it from Nevada, where Wheeler and Wheeler (1978:394) found it be- tween 6400 and 10,500 ft. They found it fre- quently in thatched and duff mounds in North Dakota (1963). Cole (1942:380) in- dicates that its nests in Utah are domed mounds of detritus in open areas usually next to a shrub at lower elevations, or of mounds of detritus in areas of moderate to dense cov- er in aspen areas. Ingham (1959, 1963) found it in mounds of detritus in juniper in southern Utah. Knowlton (1975:3) found it associated with sagebrush, rabbitbrush, and sunflowers in northern Utah. Four ants in one collection were taken from a mound of small sticks and soil next to a log in association with Formica integroides and F. obscuriventris. Ten ants in one collec- tion were taken from a mound of sticks, and sL< ants in one collection were taken from un- der a rock where F. gnava, F. neogagates, and F. perpilosa also were found. Twenty- nine ants in three collections were taken in the open. Four collections were in sagebrush 472 Great Basin Naturalist Vol. 42, No. 4 and tAvo in aspen and fir. In 71 recorded Utah habitats it was taken 24 times in mon- tane forest. In 40 recorded elevations be- tween 3000 and 9100 ft it was taken most frequently between 4000 and 5000 (25 times) and 6000 and 9000 (14 times). Beck et al. (1967:70) found it feeding on dead rodents in four instances in Utah. Formica obscuriventris Mayr F. trimcicola var. obscuriventris Mayr, 1870, Verb. Zool.- Bot. Ges. Wien 20:951. F. tnincicola obscuriventris: Rees and Grundmann 1940:8. F. tnincicola obscuriventris var. aggerans: Rees and Grundmann 1940:8. F. rufa clivia: Cole 1942:380. F. obscuriventris: Ingham 1959:80. F. obscuriventris clivia: Smith 1979:1460. Records (Map 11); BOX ELDER: Clear Crk (in Raft River Mts) (U), Mantua (KU), Promontory Pt (RG), Wellsville Mts, Willard Basin (KU). CACHE: .\nt Val- ley, Blacksmith Fk Cyn, Hyrum (KU), Logan (C42), Lo- gan Cyn, Mendon Cold Spngs, Millville, Providence, Rock Crk, Smithfield Cyn, Tony Grove (KU). GRAND: Moab (U). IRON: Cedar City (RAU). KANE: Long Val- ley Jet 11 mi W (159). RICH: Meadowville Summit 8 mi W (KU). SALT LAKE: locality unknown (RG). SAN- PETE: Majors Flats (KU). UINTAH: Whiterocks (KU). UTAH: Payson Cyn 7.3 and 10.3 mi up, Santaquin 7.7 mi E, Tibbie Fk Cyn (A). WASATCH: Trancis 18.5 mi E, Hanna 14.3 mi W (A). WEBER: Slaterville (US). Creighton (1950) and Smith (1979:1460) list two races of this species, obscuriventris and clivia, from eastern to western United States, including Utah and Colorado, nesting under logs and stones in grasslands, woods, and forests. Although both races have been recorded from Utah, the records probably are of clivia. Gregg (1963:572) lists the species between 5354 and 10,000 ft under rocks, logs, and in thatched domes in conifers, can- yons, and pinyon-juniper-oak habitats in Col- orado. Wheeler and Wheeler (1979:394) found it between 6300 and 10,000 ft in Ne- vada, and frequently in wood in North Da- kota (1963). Cole (1942:380) indicates its hab- itats in Utah as under logs and stones in sparse to moderate cover. Ingham (1959) foimd it under logs in fir, aspen, and ponde- rosa pine in southern Utah. Forty-nine ants in four collections were taken from under rocks, once under the same one with F. subnitens, once F. lasioides, and once Lasius alienus and L. niger. Eight ants in one collection were taken from a mound of small sticks and soil by a log in association with F. integroides and F. obscuripes. One ant was taken imder the same log with Camp- onotiis modoc, F. podzolica, and F. subnitens. Twenty-five ants were taken in the open. Six collections were in aspen: three in open meadows in association with fir, one with fir, and one with grass and herbs. One collection was in an association of herbs, Oregon grape, shrubs, and fir. In 30 recorded Utah habitats it was taken 18 times in montane areas. In 11 known elevations between 4217 and 8100 ft it was taken most frequently under 5000, twice over 7000. Formica obtusopilosa Emery F. sanguinea obtusopilosa Emery, 1893, Zool. Jahrb. Syst. 7:643. F. obtusopilosa: Cole 1942:377; Ingham 1963:104; Gregg 1963:610; Knowlton 1970:210, 1975:4; Allred and Cole 1979:98; Smith 1979:1450. Records (Map 12): BOX ELDER: Bear River City, Blue Crk (City) (C42), Cedar Crk (City), Cedar Mil! (K75), Curlew Jet (K70), Fielding, Garland (C42), Hansel Mts (K75), Hardup (C42), Kelton (US), Kelton Pa.ss (K70), Portage (KU), Snowville (K70), Tremonton (C42). CACHE: Cache Jet, Green Cyn (C42), Hyde Park (KU), Logan (C42), Providence (KU), Richmond (US). CAR- BON: Helper (KU). DUCHESNE: Bonita (= Boneta), Duchesne (C42) and 2 mi N (A), Fruitland (C42). EMERY: Orangeville 14.3 mi W (A). GRAND: Thomp- son (C42). IRON; Cedar Valley (159), Modena 5.3 mi W (A). JUAB: Ferno Valley (C42'). KANE: Glen Cyn City (AC). RICH: Randolph's and 10 mi SW (KU). SALT LAKE: Dry Cyn, Salt Lake City (C42). SAN JUAN: La Sal Jet 1.6 mi S (A). SANPETE: Fairview 2 mi N (A). TOOELE: Gold Hill, Skidl Valley, Stansbury Island, Timpie (C42). UINTAH: Bonanza 14 mi S (A), Dinosaur Nat Mon (Gr63), Lapoint (C42), Whiterocks (KU). UTAH: Lehi (C42), Spanish Fk Cyn (KU). WEBER: Jet U39 and U166 3.3 mi E (A). COUNTY UNKNOWN; Uinta (C42) (? = Uintah in Weber Co). Smith (1979:1450) lists this species from midwest to western United States, including Utah and Nevada, nesting under stones or in open areas with irregular mounds or craters. Gregg (1963:610) lists it from Colorado be- tween 4800 and 9500 ft under rocks and pre- dominantly in grass and sagebrush habitats, and gives a record for Utah. Cole (1966:26) found its nests in southern Nevada in soil without mounds in desert shrub habitats. Wheeler and Wheeler (1963) found it fre- quently under rocks in North Dakota. Allred and Cole (1979:98, 1971:239) found it in southern Utah and Idaho in associations of December 1982 Allred: Ants of Utah 473 ephedra-grass, sagebrush-grass, and goosefat- winterfat. Cole (1942:378) indicates that in Utah it nests in obscure craters, under stones, or in soil without craters in grassy areas. Ing- ham (1963) found it in juniper, sagebrush, and grease wood in southern Utah. Knowlton (1975:4) found it associated with rabbitbrush, sunflowers, and shadscale in northern Utah. There were 109 ants in five collections taken from small crater mounds about one inch high and six inches in diameter. Twenty ants in one collection were taken from under a rock, and 5 ants in one collection singly in an open area. One of the mound colonies contained eggs, larvae, and pupae in late July. Five collections were in sagebrush: two in association with grass and one with herbs. One collection was in a grass, juniper, and pinyon association. In 48 recorded Utah habi- tats it was taken only 4 times in montane areas. In 25 recorded elevations between 3250 and 7000 ft it was taken most frequent- ly between 4000 and 6000. Formica occulta Francoeur F. occulta Francoeur, 1973, Soc. Ent. dii Quebec, Mem. 3, p. 94; Smith 1979:1454. Records (Map 11): BOX ELDER: Willard Basin (KU). CACHE: Ant Valley, Antelope Valley, Blacksmith Fk Cyn L Fk, Franklin Basin, Tony Grove, Logan Cyn (kU). EMERY; Joes Valley (U). RICH; Randolph 10 mi SW, Sage Crk, Sage Crk Jet (KU). SEVIER; Fremont 9.7 mi N (A). SUMMIT: Wyoming brdr on U150 (A). WASATCH: Hanna 9.2 mi W (A). Smith (1979:1454) lists this species from western United States, including Utah, Colo- rado, Arizona, and Wyoming. Thirty-five ants in three collections were found under rocks, once in association with F. limata. Three collections were in sage- brush, once in association with matchbrush. In 14 recorded Utah habitats it was taken in montane areas nine times. Five recorded ele- vations were between 6260 and 9300 ft, mostly above 8000. Formica opaciventris Emery F. exsectoides var. opaciventris Emery, 1893, Zool. Jahrb. Syst. 7:643. Records: CACHE: Blacksmith Fk Cyn, Elk Valley, Frankhn Basin. Tonv Grove Lake (KU). Smith (1979:1456) lists this species from midwest to western United States, including Colorado and Wyoming, nesting in earthen or thatched mounds. Gregg (1963:601) lists it between 5160 and 10,500 ft in thatched domes from sagebrush to conifer habitats in Colorado. Formica oreas Wheeler F. oieas Wheeler, 1903, Bull. Amer. Mus. Nat. Hist. 19:643; Rees and Grundmann 1940:9; Cole 1942:379; Ingham 1959:80; Knowlton 1975:4. F. oreas comptula: Rees and Grundmann 1940:9; Cole 1942:380; Smith 1979:1461. F. oreas oreas: Smith 1979:1461. Records (Map 12); BOX ELDER: Bear River, Clear Crk, Hansel Mts (KU), Kelton (US), Kelton Pass (K75), Snowville, Willard Basin (KU). CACHE: Avon, Beaver Crk (KU), Blacksmith Fk (RG), Carter Crk (KU), Cornish (RG), Cove (C42), Franklin Basin, High Crk, Leeds Cyn (KU), Logan (C42), Logan Cvn, Tony Grove Cyn (KU). DUCHESNE: Wolf Crk (RG). KANE; Duck Crk on Ce- dar Mt (159). RICH; Allen Cyn, Sunrise Cmpgnd (KU). SALT LAKE: Big Cottonwood Cyn, Butterfield Cyn, Red Butte Cyn, Rose Cyn (U). SAN JUAN: Hole-in-the- Rock Cyn (U). UINTAH: Dry Fk rd 15 mi N U121 (A). UTAH:'Diamond Fk Cyn, Spanish Fk Cyn (KU), Thistle 14.6 and 20.4 mi E (A). WASATCH; Soldier Summit 1 mi E (A). WASHINGTON; St George (RG). COUNTY UNKNOWN: Duck Crk in Cedar Mts (RG). Creighton (1950) and Smith (1979:1461) list two races of this species from midwest to western United States, including Utah, Colo- rado, and Wyoming, nesting under logs or stones banked with detritus in open woods, meadows, or grasslands. Both races have been recorded from Utah. They may be separated by the color of the head and thorax, which in oreas are clear red, and in comptula are deep brown. Gregg (1963:577) lists this species be- tween 5200 and 10,505 ft under rocks in a variety of habitats in Colorado. Wheeler and Wheeler (1978:394) found it between 6400 and 8800 ft in Nevada, frequently in wood and commonly in duff-covered mounds in North Dakota (1963). Allred and Cole (1971:239) found it in Idaho in associations of rabbitbrush-sagebrush-grass and wild rye- grass. Cole (1942:379) indicates its habitat in Utah as under stones banked with detritus in open sunny areas. Ingham (1959) found it in mounds of detritus in southern Utah. Knowl- ton (1975:4) found it associated with rabbit- bi-ush in northern Utah. Fifty ants in two collections were taken from low mounds of sticks, both times in as- sociation with F. integroides. Twenty ants in two collections were taken singly in open 474 Great Basin Naturalist Vol. 42, No. 4 areas. Four collections were in sagebrush: one in association with herbs; one grass and rabbitbrush; one grass, clover, and Russian thistle; and one grass, aspen, and pine. In 36 recorded Utah habitats it was taken 23 times in montane forest. In 15 recorded elevations between 2760 and 9300 ft it was taken most frequently between 4000 and 5000 ft. Formica pallidefulva Emery F. palUde-fitlva nitidiventris Emery, 1893, Zool. Jahrb. Syst. 7:645; Rees and Gnindmann 1940:10. F. pallidefulva nitiventris: Ingham 1959:85. F. pallidefulva: Knowlton 1975:4. Records (Map 12): BOX ELDER: Wildcat Hills (K75). GRAND: La Sal Mts (KU). JUAB: Nebo Loop rd 9.9 mi S Santaquin Cyn (A). SALT LAKE: Parleys Cvn (RG). UINTAH: Red Cloud Loop rd 9.2 mi W U44 (A). WASATCH: Francis 8.1 mi E (A). WASHINGTON: Zion Nat Park (159). Creighton (1950) and Smith (1979:1451) list two forms of this species from eastern to western United States, including Colorado and Wyoming. The Utah subspecies probably is nitidiventris, which can be distinguished from pallidefulva by the clear golden yellow color of pallidefulva as contrasted with the yellowish to blackish brown of nitidiventris. Gregg (1963:627) lists this species between 3500 and 8000 ft under rocks in a variety of habitats in Colorado. Ingham (1959) found it under rocks in ash, oak, poplar, and poison ivy in southern Utah. Knowlton (1975:4) found it associated with sagebrush in north- em Utah. Fifty ants in two collections were taken from under rocks, once under the same one with F. neoclara, and once with F. argentea and Pheidole desertorum. Three ants in one collection were taken singly in an open area. One collection was in sagebrush, one cotton- woods, and one a grassy meadow. In seven recorded Utah habitats five were in montane areas. One recorded elevation is 4500 ft. Formica pergandei Emery F. pergandei Emery, 1893, Zool. Jahrb. Syst. 7:643. Record: UINTAH: Whiterocks Cyn (KU). Smith (1979:1464) lists this species from eastern to western United States, including Colorado, frequently associated with F. fusca and F. pallidefulva. Formica perpilosa Wheeler F. fusca subpoUUi var. perpilosa Wheeler, 1902, Mem. Revist. Soc. Sci. Ant. Alzate 17:141. F. perpilosa: Rees and Grundmann 1940:8; Cole 1942:378; Grundmann 1958:166; Ingham 1959:77; Beck et al. 1967:70. Records (Map 13): EMERY: San Rafael River (BAD). KANE: Cottonwood Cyn (BAD), Glendale, Kanab, Or- derville (C42). MILLARD: Sutherland (US). PIUTE: Marysvale 4.1 mi S (A). SALT LAKE: Salt Lake City (RG). TOOELE: Stansbury Island (RG). UINTAH: Jen- sen (U). WASHINGTON: Grafton (159), Hurricane (RG), Rockville (159), Santa Clara (C42), Springdale (159), St George (C42), Washington (KU). WAYNE: Pleasant Crk (BAD). Smith (1979:1450) lists this as a mid- western and western species, including Colo- rado and Wyoming, which usually constructs crater nests in grasslands and open fields. Hunt and Snelling (1975:23) list it from Ari- zona. Gregg (1963:611) lists it between 3500 and 6000 ft under wood and in dome nests in a variety of habitats in Colorado. La Rivers (1968:10) lists it from Nevada. Cole (1942:378) states that in Utah its nests are ob- scure craters or low domes around the roots of trees and shrubs, particularly in irrigated areas and dry river beds. Grundmann (1958:166) indicates that its nests in Utah are obscure and difficult to find, consisting of mounds around the bases of shrubs and trees along stream banks. Ingham (1959, 1963) found it in southern Utah under wood, in small half-moon-shaped mounds, and in clumps of grass in willow, tamarix, poplar, and alkali flats. One ant was collected from under a rock in sagebrush. Ants of F. gnava, F. neogagates, F. obtusopilosa, and F. perpilosa were under the same rock. In 18 recorded Utah habitats only one was in a montane area. Twenty-one collections at known elevations were about equally distributed between 2625 and 6600 ft. Beck et al. (1967:70) found it feeding on dead rodents in three instances in Utah. Formica podzolica Francoeur F. podzolica Francoeur, 1973, Ent. Soc. Quebec, Mem. 3:167. Records (Map 13): BEAVER: Beaver (F). BOX EL- DER: Cedar Hill (US), Clear Crk (in Raft River Mts) (U), Snowville (US), Willard Pk (KU). CACHE: Ant Val- ley, Bear River Mts, Blacksmith Fk Cyn, Franklin Basin, Green Cyn, Hodges Cyn, Hyrum, Logan (KU), Logan Cvn (US), Millville, Providence, Tony Grove (KU), Wellsville Mts (US). CARBON: Clear Crk Cyn (F), December 1982 Allred: Ants of Utah 475 Scofield 5.2 mi SW, Scofield Res 4 mi S, Wellington 12 mi NE (A). DAVIS: Farmington (KU), Farmington Cyn (US). IRON: Upper Bear Crk, Cedar City (RAU) and 19 mi E (A), Summit Crk (RAU). JUAB: Lynndyl 11.5 mi N (A), Toms Crk (in Deep Crk Mts) (U). RICH: Allen Cyn (F), MeadowviUe, Monte Cristo, Pickleville (KU). SAN JUAN: Elk Ridge, Kigalia Ranger Sta (F). SUMMIT: Mirror Uke 11.6 and 17.3 mi N (A). TOOELE: Dugway 11.9 mi E (A), Willow Cyn (F). UINTAH: Dry Fk rd 15 mi N U121, Red Cloud Loop rd 4.2 and 9.2 mi W U44 (A), Whiterocks (F). UTAH: Mt Nebo (F), Payson Cyn 12.3 mi up, Santaquin Cyn 4.5, 6.7, 7.1 and 7.7 mi up (A), Thistle (F), Tibbie Fk Cyn, Tibbie Fk Lake 2.3 mi W (A). WASATCH: Francis 8.1 and 18.5 mi E, Hanna 14.3 mi W, Midway 3.7 mi NW, Soldier Summit 1 mi E and 10.1 mi N (A). WEBER: Beaver Crk (KU). COUN- TY UNKNOWN: Beaver Head (KU), Chalk Crk (F) (in either Millard or Summit Co). Smith (1979:1455) lists this species from eastern to western United States, including Arizona, nesting in soil mounds in montane forest. There were 123 ants in 10 collections taken from under rocks, once under the same one with F. gnava, once F. fusca, once F. moki, once F. limata, and once F. fusca and Solenopsis molesta. There were 171 ants in 6 collections taken from ground burrows, only 2 of which had low mounds. Twenty-three ants in 4 collections were taken from under logs, once under the same log with F. fusca and F. gnava, once Camponotus modoc, and once C. modoc, F. obscuriventris, and F. sub- nitens. Thirty ants in one collection were taken from inside a log. Forty-nine ants in 6 collections were taken singly in open areas. Immatures were found in a burrow in late June and under a rock in early August. Six- teen collections were from aspen: 4 in associ- ation with grass, herbs, and shrubs; three sagebrush and conifers; one grass; one herbs in an open meadow; one chokecherry; one chokecherry and fir; one maple, oak, and fir; and one pine. Two collections were in grass and herbs, one juniper and pinyon, two fir, one oak, and one sagebrush and herbs. In 62 recorded Utah habitats 41 were in montane forest. In 15 recorded elevations between 4302 and 9300 ft it was taken 11 times under 6000. In one instance, when a cover rock was removed to expose the ants, they ran rapidly and tried to escape. In another case, when the burrow was excavated, the ants scurried around but did not leave the area. Many of them worked furiously to remove the few ex- posed pupae. Formica puberula Emery F. sanguinea puberula Emery, 1893, Zool. Jahrb. Syst. 7:643; Rees and Grundmann 1940:8; Cole 1942:378; Knowlton 1970:210. F. puberula: Creighton 1950:468; Gregg 1963:614; Knowlton 1975:4. Records (Map 13): BOX ELDER: SnowviUe (K75). GRAND: Warner Ranger Sta (Gr63). PIUTE: Fish Lake Jet 1 mi S (U). SAN JUAN: La Sal Mts (RG). SANPETE: Ephraim Cyn (KU), Wales (RG). TOOELE: Stockton (RG). UINTAH: Paradise Park 11 mi S (U). Smith (1979:1465) lists this species from midwest and western United States, including Colorado and Wyoming, where it frequently associates with as many as 12 species of For- mica. Gregg (1963:614) lists it between 5100 and 10,000 ft under rocks and logs pre- dominantly in conifer habitats in Colorado, and gives a record for Utah. La Rivers (1968:10) lists it from Nevada, where Wheel- er and Wheeler (1978:396) found it between 6400 and 8200 ft. Cole (1942:378) indicates that it nests under stones in Utah. In seven recorded Utah habitats it was found three times in montane areas. Three known eleva- tions are 4544, 5069, and 8000 ft. Formica querquetulana Kennedy and Dennis F. querquetulana Kennedy and Dennis, 1937, Ann. Ent. Soc. Amer. 30:536. Record: CACHE: Leeds Cyn (KU). Smith (1979:1463) lists this species from eastern United States. Its occurrence in Utah is questionable. Formica rasilis Wheeler F. microgyria var. rasilis Wheeler, 1903, Bull. Amer. Mus. Nat. Hist. 19:648. F. microgyria rasilis: Rees and Grundmann 1940:9; Cole 1942:381. F. rasilis: Creighton 1950:506; Gregg 1963; Knowlton 1975:5; Smith 1979:1463. Records (Map 13): BOX ELDER: Snowville (K75). CACHE: Bear River Mts (KU). GARFIELD: Bryce Cyn Nat Park (Cr). GRAND: Warner Ranger Sta (Gr63). SALT LAKE: Butterfield Cyn, Red Butte Cyn (U). SAN JUAN: La Sal Mts (RG). Smith (1979:1463) lists this species from western United States, including Utah and Colorado, nesting under stones in open areas. Gregg (1963) lists it between 5400 and 11,542 ft under rocks and logs and in thatched nests predominantly in conifer habi- tats in Colorado, and gives a record for Utah. 476 Great Basin Naturalist Vol. 42, No. 4 La Rivers (1968:10) lists it from Nevada. Cole (1942:381) indicates its habitat in Utah as un- der stones frequently banked with detritus. In six recorded Utah habitats five were in mon- tane areas. Six recorded elevations were be- tween 4544 and 8000 ft. Formica subintegra Emery F. sanguinea rubicunda var. subintegra Emery, 1893, Zool. Jahrb. Syst. 7:643. Records: BOX ELDER: Hansel Mts (KU). RICH; Monte Cristo (KU). SANPETE: Ephraim Cyn (KU). Smith (1979:1465) lists this species from eastern to midwestern United States. No in- termountain state is listed, and its occurrence in Utah is questionable. Formica subnitens Creighton F. rufa subnitens Creighton, 1940, Amer. Mus. Nat. Hist. Nov. 1055:7. F. subnitens: Knowlton 1970:210, 1975:5. F. subaenescens: Knowlton 1975:5. Records (Map 14): BOX ELDER: Hardup, Snowville (K70) and 9 mi W (US). CACHE: Elk Valley, Leeds Cyn, Logan Cyn, Rock Crk (in Blacksmith Fk Cyn), Tony Grove Cyn (KU). IRON: Upper Bear Crk (RAU). RICH: Sage Crk Jet 5.1 mi W (A). UINTAH: Drv Fk rd 15 mi N U121, Red Cloud Loop rd 4.2 mi W U44 (A). UTAH; Halls Fk rd 8.8 mi N Hobble Crk rd, Payson Cyn 10.3 mi up, Santaquin Cyn 7.7 mi up (A). WASATCH: Midway 5.7 and 11.6 mi W, Soldier Sum- mit 3.3 mi N, Strawberry Res 4 mi S (A). Smith (1979:1461) lists this species from midwest to western United States, including Colorado and Wyoming, nesting under stones or in mounds of thatch. Gregg (1963:581) lists it at 6100 ft in thatched nests in pinyon-juni- per in Colorado. Wheeler and Wheeler (1963) found it in thatched moimds in North Dakota. Seventy-five ants in three collections were taken from mounds of sticks, once next to a boulder and once at the base of a stump. One ant was taken from under the same log as F. obscuriventris, F. podzolica, and Camponotus niodoc. Eleven ants in two collections were taken from under rocks, once under the same rock with F. obscuriventris. Thirteen ants in four collections were taken singly in open areas. Eggs were found in one mound in late July. Five collections were in aspen: one in association with grass and herbs; one grass, herbs, shrubs, and fir; one grass, sagebrush, and pine; one chokecherry; and one fir. Two collections were in grass, herbs, and sage- brush; one legumes and sagebrush; one matchbrush and sagebrush; and one oak and fir. In 19 recorded Utah habitats it was taken 15 times in montane areas. One recorded ele- vation was 4544 ft. Ants on a mound of thatched soil were highly defensive, rearing back in a position of defense to await an invader. Formica subnuda Emery F. sanguinea rubicunda var. subnuda Emery, 1895, Zool. Jahrb. Syst. 8:335. F. sanguinea subnuda: Rees and Gnmdmann 1940:8; Cole 1942:378; Hayward 1945:120; Creighton 1950:469. F. subnuda: Ingham 1959:79; Gregg 1963:620. Records (Map 14); BOX ELDER: Snowville (C42). CACHE: Tonv Grove (KU). CARBON: Scofield 4 mi S (A). GARFIELD: Boulder Mt (U), Bryce Cvn Nat Park (WU). GRAND; Warner Ranger Sta ('Gr63).' IRON: Ce- dar Breaks Nat Mon (159), Cedar City 14 mi E (A). KANE: Cedar City 24.3 mi E (A). SALT LAKE: Big Cottonwood Cvn, Red Butte Cyn (U). SAN JUAN; La Sal Mts (C42), Monticello 5 mi W (U). SANPETE: Or- angeville 23.9 mi W (A), Wales (C42). SUMMIT: Kamas 11 mi E (U), Kamas 21 and 28.5 mi E (A), Soapstone Cyn (RG). TOOELE; Grantsville, Stockton, Tooele (C42). UINTAH: East Pk Res rd 2 mi W U44 (A), Trial Lake (U). UTAH; Aspen Grove, Provo Cyn (U), Silver Lake Flat (A). WASATCH: Horse Crk (C42), Midway 11.6 mi W, Soldier Summit 7.9 mi N (A). WAYNE: Capitol Reef Nat Park (U), Henrys Fk Basin (RG). COUNTY UN- KNOWN: Pallisade Park (in Ashley Nat Forest) (RG). Smith (1979:1465) lists this species from eastern to western United States, including Colorado and Arizona, frequently associated with F. altipetens, F. fiisca, F. neorufibarbis, and F. siibpolita. Gregg (1963:620) lists it be- tween 5000 and 13,000 ft under rocks and logs and in thatched nests in a variety of hab- itats in Colorado, and gives a record for Utah. La Rivers (1968:11) lists it from Ne- vada, where Wheeler and Wheeler (1978:396) found it between 8200 and 11,000 ft. They found it frequently in wood, also common under rocks and in earthen mounds in North Dakota (1963). Cole (1942:378) in- dicates it nests under stones and logs in Utah. Ingham (1959) found it under stones and logs in fir, spruce, aspen, and pine in southern Utah. There were 244 ants in nine collections found under logs, once under the same one with F. fusca, once F. gnava, and once F. manni. Fifteen ants in one collection were December 1982 Allred: Ants of Utah 477 found inside a decaying log. These tried to escape when disturbed, crawling under de- bris. One ant next to the log had a struggling carabid beetle in its jaws, carrying it toward the log. Sixteen ants in one collection were taken singly in an open area. Eggs were found under one log in mid-July. Ants associ- ated with these eggs held firmly onto the sub- strate and reared back in a biting position when disturbed. Eight collections were in as- pen: one in association with grass; one grass, herbs, and pine; two conifers; and one choke- cherry. Three collections were in conifers. In 32 recorded Utah habitats it was taken 26 times in montane forest. In 15 recorded ele- vations between 4304 and 10,500 ft it was taken more frequently above 7000. Formica subpolita Mayr F. fusca var. subpolita Mayr, 1886, Verh. Zool.-Bot. Ges. Wien 36:426. F. subpolita cemponoticeps: Rees and Grundmann 1940:10; Cole 1942:384; Ingham 1959:84; Knowl- ton 1970:210, 1975:5. F. subpolita: Cole 1942:383; Ingham 1959:83. F. subpolita ficticia: Creighton 1950:542. Records (Map 14): BOX ELDER: Bear River City, Blue Crk (City), Cosmo (? = Kosmo), Hardup, Kelton, Locomotive Spngs, Park Valley, Penrose, Rosette, Snow- ville, WiUard (C42). CACHE:' Logan, Logan Cyn, Tren- ton (C42). EMERY: San Rafael Swell (US). GRAND: Moab (RG). JUAB: Diamond Cyn, Nephi (C42). MIL- LARD: Delta (C42), Swasey Spngs (RG). SALT LAKE: Ft Douglas (C42). SAN JUAN: Blanding (C42), La Sal Mts (U). TOOELE: Clover, Fisher Pass, Flux, Grants- ville, losepa, Orrs Ranch, Stansbury Island (C42). UTAH: Provo (C42). WASHINGTON: Hurricane (RG), Pine Valley (City) (159), St George (C42). WEBER: Og- den (C42),' COUNTY UNKNOWN: Rosebud, Showell, Westpoint (C42). Smith (1979:1455) lists this species from western United States, including Nevada and Idaho, nesting in mounds or craters in semi- desert areas. Gregg (1963:541) lists it from Colorado between 5300 and 7800 ft. Wheeler and Wheeler (1978:396) found it between 6000 and 10,800 ft in Nevada. Allred and Cole (1971:239) found it in Idaho in associ- ations of a variety of shrubs. Cole (1966:26) found its nests in southern Nevada under stones commonly in pinyon-junipep; also in grassy areas in Utah (1942:384). Ingham (1959, 1963) found it under stones in juniper and sagebrush in southern Utah. Knowlton (1975:5) found it in northern Utah. In 38 recorded Utah habitats it was taken only 3 times in montane areas. In 28 re- corded elevations between 2700 and 9000 ft it was taken most frequently between 3000 and 4000, twice over 7000. Formica subsericea Say F. subsericea Say, 1836, Boston J. Nat. Hist. 1:289. Records: CACHE: Logan Cyn, WellsviUe (US). SEVIER: Richfield (US). Smith (1979:1455) lists this species from eastern to midwestern United States, but does not include an intermountain one. Wheeler and Wheeler (1978:396) found it between 7000 and 11,500 ft in Nevada, and frequently in earthen mounds, under rocks, and in and under wood in North Dakota (1977). Utah records of this species were taken at 4495 and 5340 ft. According to Snelling (pers. comm.) these Utah records are probably based on misidentifications. Formica transrnontanis Francoeur F. transmontanis Francoeur, 1973, Soc. Ent. du Quebec, Mem. 3:35. Record: SAN JUAN: Monticello (KU). Smith (1979:1455) lists this species from western United States, including Idaho. Formica wheeleri Creighton F. wheeleri Creighton, 1935, Amer. Mus. Nat. Hist. Nov. 773:1; Rees and Grundmann 1940:10; Cole 1942:378; Creighton 1950:472; Grundmann 19,58:166; Smith 1979:1466. Records: GRAND: Warner Ranger Sta (RG). SAN JUAN: Blue Mts (C42) (? = Abajo Mts). Smith (1979:1466) lists this species from midwest and western United States, including Utah, Colorado, and Arizona, associated with F. altipetens, F. fusca, F. lasioides, F. neo- gagates, and F. neorufibarbis. Gregg (1963:623) lists it between 5500 and 9500 ft under rocks in several habitats in Colorado. Wheeler and Wheeler (1963) found it under rocks and in earthen mounds in North Da- kota. Cole (1942:378) states that it nests un- der stones on open hillsides in aspen forests in Utah, where it takes F. lasioides as a slave species (Grundmann 1958:166). Formica whymperi Wheeler F. adamsi var. alpina Wheeler, 1909, J. New York Ent. Soc. 17:85. 478 Great Basin Naturalist Vol. 42, No. 4 F. whymperi alpina: Creighton 1950:509; Gregg 1963:593; Smith 1979:1463. Record: GRAND: Warner Ranger Sta (Gr63). Creighton (1950) and Smith (1979) hst four races of this species from midwest to western United States, including Utah, Colorado, and Idaho, nesting under stones or logs in forest areas, where it associates with F. neoclara and F. neorufibarbis. The subspecies in Utah is likely alpina. Gregg (1963:593) lists it be- tween 8500 and 12,500 ft under rocks and logs and in thatched nests in conifer habitats in Colorado, and gives a record for Utah. All- red and Cole (1971:239) found it in associ- ations of rabbitbrush-sagebrush-grass-winter- fat in Idaho. Formica xerophila M. R. Smith F. nwki xerophila Smith, 1939, Ann. Ent. Soc. Amer. .32:5a3. F. moki: Rees & Grundmann 1940:10; Ingham 1959:84. F. xerophila: Francoeur 1973:262; Smith 1979:1455. Records (Map 14): BEAVER: Milford (RG). DAVIS: Farmington Cyn (KU). SALT LAKE: Big Cottonwood Cyn, Mt Olympus (U), Parleys Cyn (Cr). SAN JUAN: Blanding, Bluff (RG). UINTAH: Red Cloud Loop rd 4.2 mi W U44 (A). WASHINGTON: Big Plains, Central, Zion Nat Park (159). Smith (1979:1455) lists this species from western United States, including Utah and Arizona. In 12 recorded Utah habitats 5 were in montane areas. Fourteen recorded eleva- tions were between 4000 and 7000 ft, 6 un- der 5000. Formicoxenus chamberlini (Wheeler) Symmyrmica chamberlini Wheeler, 1904, Bull. Amer. Mus. Nat. Hist. 20:5; Rees and Grundmann 1940:6; Cole 1942:370; Creighton 1950:281; Smith 1979:1.398. Records: SALT LAKE: Salt Lake City (RG). SAN JUAN: Blanding 8 mi N (U). Smith (1979:1398) lists this species as Utah and Oregon, nesting with Manica mutica. Hypoponera opaciceps (Mayr) Ponera opaciceps Mayr, 1887, Verh. Zool.-Bot. Ges. Wien 37:536; Grundmann 1958:161; Ingham 1963:39. Records: SALT LAKE: Salt Lake City (U). WASH- INGTON: St George (159). According to Smith (1979:1343), this is pri- marily an eastern species ranging westward to Arizona and Colorado. Gregg (1963:284) found it between 5354 and 5400 ft in cotton- wood-willow habitats in Colorado. La Rivers (1968:2) listed it from Nevada. Grundmann (1958:161) indicates that in Utah it ranges up to 4000 ft in dry desertlike areas where there is moisture, and nests under stones among willows. Four known elevations in Utah are from 2700 to 4453 ft. Hypoponera opacior (Forel) Ponera trigona var. opacior Forel, 1893, Trans. Ent. Soc. London, p. 363; Cole 1942:359. Records: SALT LAKE: Salt Lake City (U). UTAH: Springville (C42). Smith (1979:1343) lists this species from eastern to western United States, including Colorado. Hunt and Snelling (1975:20) list it from Arizona. Gregg (1963:286) lists it be- tween 4000 and 5400 ft under rocks in grassy habitats in Colorado. One Utah collection was taken at 4253 ft. Iridomyrmex humilis (Mayr) Hypoclinea humilis Mayr, 1868, Soc. Nat. Modena, Ann. 3:164. Records: GRAND: Moab (U). SAN JUAN: Goulding Trading Post (U). Smith (1979:1418) lists this species from eastern to western United States, including Arizona, nesting in soil, rotting wood, or debris. La Rivers (1968:6) lists it from Ne- vada. Elevation of two recorded Utah collec- tions was 4000 ft. Iridomyrmex pruinosus (Andre) Tapinoma anale Andre, 1893, Rev. Ent. de France 12:148. /. pruinosus: Rees and Grundmann 1940:7; Cole 1942:.373; Allred and Cole 1979:98. /. analis: Rees and Grundmann 1940:6. /. pruinosus analis: Rees and Grundmann 1940:7; Cole 1942:373; Grundmann 1958:165; Ingham 1959:62; Beck et al. 1967:70. /. pruinosus testaceus: Cole 1942:373. Records (Map 15): BOX ELDER: Bovine (C42), Brig- ham (A), Lucin, Park Valley (City) (C42). CACHE: Green Cyn (KU), Logan Cyn (C42), Tony Grove (KU). DUCHESNE: Roosevelt (C42). EMERY: Greenriver (US), Wellington 46 mi S (A). GRAND: Dewey (U), Moab (C42). JUAB: Chicken Crk Res (KU). KANE: Castle Rock (U), Coral Pink Sand Dunes (159), Glen Cyn City (AC), Kanab (C42). MILLARD: Deseret (C42). SALT LAKE: Big Cottonwood Cyn (C42), Butterfield Cyn (U), S Dry Cyn, Ft Douglas (C42), Mt Olympus (U), December 1982 Allred: Ants of Utah 479 Parleys Cyii (C42). SAN JUAN: Bluff (BAD), Johns Cyn (RG), La Sal (C42), Monticello (U), Montezuma Crk (BAD), Monument Valley (KU). SEVIER: Salina Cyn nr Fremont Jet (U). TOOELE: Clover, losepa (C42). UIN- TAH: Gusher (C42). UTAH: Aspen Grove (BY), Pelican Pt (RG). WASHINGTON: Harrisburg (US), Harrisburg Jet (159), Hurricane (US), La Verkin, Pintura, Rockville, Vevo, Virgin Citv (159), Zion Nat Park (C42). WAYNE: Friiita 5 mi SE (U), Hanksville 17 mi S (KU). COUNTY UNKNOWN: Valley Jet, Willow Spngs (C42). Snelling (pers. comm.) indicates that this is a Forelius, but for the present I am retaining it as indicated. Creighton (1950) and Smith (1979:1419) indicate two races of this species as midwest and western in the United States, inckiding Idaho, where nests are imder objects or in craterhke moimds in open areas. The Utah population, probably the subspecies onalis, can be separated from pruinosus by the dense pubescence on the head and thorax of prui- nosus that partially obscures its rough sur- face, whereas on analis the pubescence is di- lute and reveals the shining surface beneath. Hunt and Snelling (1975:22) list it from Ari- zona. Gregg (1963:436) lists it from Colorado between 3500 and 6300 ft under rocks in a variety of habitats, predominantly pinyon- juniper and grass. Cole (1966:18) states that in southern Nevada it nests imder stones, at the base of plants, and in mounds in open areas in a variety of desert shrub habitats. He found it in Utah in craterlike nests in sandy soil (1942:373). Wheeler and Wheeler (1963) found it in soil craters in North Dakota. Gnmdmann (1958:165) indicates that nests of these ants in Utah are difficult to locate, and occur most commonly in grass habitats near desert mountains. Ingham (1959, 1963) found it imder stones and in small craters in creo- sote bush, sagebRish, pinyon-juniper, galleta- grass, rabbitbrush, winterfat, Joshua trees, four-wing saltbush, and shadscale in southern Utah. Allred and Cole (1979:98) found it in southern Utah in a variety of desert shrub types, most commonly in sagebrush. Sixty ants in two collections were taken from small mounds with a central top open- ing. These ants are rapid runners. One collec- tion was in halogeton and one in sunflowers, sagebrush, and rabbitbrush. In 49 reported habitat localities this species was taken 10 times in montane areas. In 45 recorded eleva- tions between 2500 and 8100 ft it was taken 29 times between 3000 and 5000, 14 times between 5000 and 8000. Beck et al. (1967:70) found it feeding on dead rodents in three in- stances in Utah. Lasius alienus (Foerster) Formica aliena Foerster, 1850, Hymenop. Studien (Ernst Ter Meer Publ. Aachen) L.36. L. niger alienus var. americantis: Rees and Grundmann 1940:7. L. niger var. auiericanus: Rees and Grundmann 1940:7; Cole 1942:374; Knowlton 1970:210. L. alienus americanus: Grundmann 1958:166; Ingham 1959:72; Knowlton 1970:210, 1975:5. L. alienus: Beck et al. 1967:70. Records (Map 15): BOX ELDER: Kelton, Kelton Pass (K70), Snowville (C42), Willard Basin (KU). CACHE: Ant Valley, Beaver Crk (KU), Blacksmith Fk Cyn, Cow- lev Cyn (C42), Franklin Basin, W Hodges Cyn (KU), Lo- gan Cyn (C42), Tony Grove Lake (KU). DUCHESNE: Duchesne (C42). EMERY: Hideout Cyn nr Green River (U). GARFIELD: Boulder Mt, Henry Mts (G58), Osiris (U). GRAND: Thompson (KU). IRON: Coal Crk Cyn (159). KANE: Kanab and 20 mi N (C42). MILLARD: lo- cality unknown (RG). RICH: Allen Cyn (KU), Randolph 2.3 mi N, Sage Crk Jet 5.1 mi W (A). SALT LAKE: Alta, Big Cottonwood Cyn, Butterfield Cyn, City Crk Cyn, Holladav, Lake Blanche, Mill Crk Cyn (C42), Mt Olympus, Red Butte Cyn (U), Salt Lake City (C42). SAN JU'aN: .-Vbajo Mts (G58), La Sal (C42), Monticello 7.6 mi W (A), Nat Bridges Nat Mon (U). SANPETE: Fairview Summit (KU). SUMMIT: Mirror Lake 6.4 mi N, Wyo- ming brdr on U150 (A). TOOELE: Clover, Fisher Pass, Tooele (C42). UINTAH: Bonanza 25 mi S, Dry Fk rd 13.4 mi N U121 (A), Gusher (C42). UTAH: American Fk Cvn, Provo (C42), Silver Lake Flat (A). WASATCH: Francis 18.5 mi E, Hanna 14.3 mi W (A). WASHING- TON: Kolob (159). Pine Valley (City), (BAD), Zion Nat Park (C42). Smith (1979:1435) indicates this as an east- ern to northwestern United States species, in- cluding Idaho and southern Arizona. It shows a preference for well-shaded woodlands, where it nests under stones and in rotting logs, only occasionally found in the open. Gregg (1963:456) lists it from Colorado be- tween 3500 and 10,400 ft under rocks in a great variety of habitats, predominantly pin- yon-juniper and grass. La Rivers (1968:8) lists it from Nevada, where Wheeler and Wheeler (1978:393) found it between 6400 and 9700 ft. They found it in wood and under bark in North Dakota (1963). Cole (1942:374) states that its habitat in Utah is under stones in open and grassy areas, some colonies at 6500 ft. In Utah it nests under stones on rocky, ex- posed mountain slopes up to 6000 ft (Gnmd- mann 1958:166). Ingham (1959) found it in southern Utah under rocks and logs in oak and aspen. 480 Great Basin Naturalist Vol. 42, No. 4 There were 155 ants in five collections taken from under rocks. In one instance Tapi- noma sessile was found under the same rock, and once alienus was under the same rock with Formica obscuriventris and Lasius niger. Sixty-nine ants in three collections were taken from under logs. In one case F. fusca was under the same log. One ant was taken from a burrow in the open. Four collections were in aspen: one in association with grass, herbs, and conifers and one with fir. Four collections were in sagebrush: two in associ- ation with grass and herbs and one with snowberry. One collection was in juniper and pinyon and one in conifers. In 56 recorded Utah habitats it was found 32 times in mon- tane forest. In 32 elevational records it was about equally distributed between 4000 and 9300 ft, slightly more common between 4000 and 6000. Beck et al. (1967:70) found it feed- ing on dead rodents in one instance in Utah. Lasius crypticus Wilson L. crypticus Wilson, 1955, Bull. Mus. Comp. Zool. 113:1 (»; Beck et al. 1967:70; Smith 1979:1436. Records (Map 15): BOX ELDER: Brigham (KU). CACHE: Logan (KU). CARBON: Myton rd 15 mi E US6 (A). GARFIELD: Henry Mts (Wi). IRON: Cedar City 14 mi E (A). KANE: Long Valley Jet (Wi). RICH: Randolph 2.3 mi N (A), Woodruff (BAD). SAN JUAN: Geyser Pass, Mexican Water (BAD), Monticello 2 mi W (A). SANPETE: Orangeville 19.5 mi W (A). SEVIER: Sa- lina Cyn nr Fremont Jet (U). UINTAH: Vernal 15 mi N (A). UTAH: Prove, Thistle 20.4 mi E (A), Wanrhodes Cyn (KU). WASATCH: Hanna 9.2 mi W (A). WAYNE: Pleasant Crk (BAD). Smith (1979:1436) lists this species from midwest to western United States, including Utah and Idaho, nesting under stones or in craters. Cole (1966:20) found its nests under stones in open areas in pinyon-juniper in southern Nevada. AUred and Cole (1971:239) found it in Idaho in a variety of shrub types. Wheeler and Wheeler (1963) found it fre- quently under rocks, also commonly in soil craters in North Dakota. Seventy-nine ants in seven collections were taken from under rocks. It was under the same rock with L. humilis, and once with Myrmica americana. Fifteen ants in one col- lection were taken from a small crater mound, and 11 in two collections singly in open areas. Five collections were in sage- brush: one in association with grass; one grass, herbs, and rabbitbrush; one herbs and juniper; and one grass, legumes, shrubs, juni- per, and pinyon. One collection was in grass, one grass and oak, one pine, and one a culti- vated area. In 19 recorded Utah habitats it was taken 9 times in montane forest. Four re- corded elevations were between 4307 and 10,750 ft. Beck et al. (1967:70) found it feed- ing on dead rodents in four instances in Utah. Lasius fallax Wilson L. fallax Wilson, 1955, Bull. Mus. Comp. Zool. 113:130; Smith 1979:14.37. Records (Map 15): CACHE: Bear River Mts, Logan Cyn (KU). GRAND: Warner Ranger Sta (Wi). SAN JUAN: Blue Mts (Wi) (? = Abajo Mts). UINTAH: Deep Crk (Wi), Red Cloud Loop rd 4.2 mi W U44 (A). WASATCH: Soldier Summit 3.3 mi N (A). COUNTY UNKNOWN: Bassets Spngs (Uinta Mts) (Wi). Smith (1979:1437) lists this species from the western United States, including Utah, Colorado, Arizona, Idaho, and Wyoming, nesting under stones in forest clearings. Fifty ants in two collections were taken from under rocks in sagebrush: one in associ- ation with grass and one with matchbrush. Eight collections whose recorded Utah habi- tats were known were in montane forest. One recorded elevation is 9750 ft. Lasius humilis Wheeler L. humilis Wheeler, 1917, Proc. Amer. Acad. Arts, Sci. 52:528. Records: IRON: Cedar City 14 mi E (A). JUAB: Iba- pah (U). KANE: Cedar City 24.3 mi E (A). UINTAH: Bonanza (KU). Smith (1979:1438) lists this species from the western United States, including Colo- rado and Nevada, nesting under stones in meadows and open woods. Gregg (1963:475) lists it between 5154 and 7000 ft under rocks in meadows in Colorado. Forty-five ants in two collections were found under logs, one in grass, herbs, aspen, and pine, and one in pine. Ants of L. sub- umbratus were under the same log in one in- stance, and L. crypticus in the other. In two of four recorded Utah habitats it was found in montane forest. Two known elevations are 5288 and 5456 ft. Lasius nearcticus Wheeler L. flavus nearcticus Wheeler, 1906, Psyche 13:38; Rees and Grundmann 1940:7; Cole 1942:374; Grund- mann 1958:166. December 1982 Allred: Ants of Utah 481 L. flavus ckiripennis: Cole 1942:375. L. flavus inicrops: Ingham 1959:74. Records (Map 17): CACHE: Logan (KU). DU- CHESNE: Paradise Park 11 mi W (U). PIUTE: Fish Lake Jet 1 mi S (U). SALT LAKE: Big Cottonwood Cyn (U), Butterfield Cyn (RG), Red Butte Cvn (U). SAN JUAN: Bluff, La Sal Mts (RG). UINTAH: Bonanza (KU). WASHINGTON: Veyo, Zion Nat Park (159). Smith (1979:1438) lists this species from eastern to western United States, including Colorado and Wyoming, where it nests under rocks or fallen logs in moist woodlands. Himt and Snelling (1975:22) list it from Arizona. Gregg (1963:471) lists it between 6250 and 9000 ft under rocks in conifers, pinyon-juni- per, and oak habitats in Colorado. La Rivers (1968:8) lists it from Nevada, where Wheeler and Wheeler (1978:393) found it between 6200 and 10,400 ft. They found it frequently under rocks in North Dakota (1963). Cole (1942:374) indicates its habitat in Utah as un- der stones. In Utah it nests under stones or dead wood among cottonwoods (Grundmann 1958:166). Ingham (1959) found it under stones in ash, oak, poplar, and poison ivy in southern Utah. In 10 recorded Utah habitats it occurred only 4 times in montane forest. In 10 eleva- tion records between 4320 and 8000 ft it oc- curred 7 times under 5000, twice over 8000. Lasius niger (Linnaeus) Formica niger Linnaeus, 1758, Syst. Nat. Ed. 10, 1:580. L. niger neoniger: Rees and Grundmann 1940:7; Cole 1942:374; Hayward 1945:120; Grundmann 1958:166; Ingham 1959:72. L. niger: Beck et al. 1967:70; Smith 1979:1436. Records (Map 16): BOX ELDER: Kelton (US), Lucin (BAD), Snowville, Wellsville Mts, WiUard Basin (KU). CACHE: Ant Valley (KU), Blacksmith Fk Cyn, Cowley Cyn (RG), Franklin Basin (KU), High Crk (US), W. Hodges Cyn (KU), Logan (RG), Logan Cyn, Tony Grove Cyn (KU). EMERY: Hideout Cyn nr Green River (U). GARFIELD: Escalante 20 mi E (U), Boulder Mt (G58), Osiris, Trachyte Ranch (U). GRAND: Moab 12 mi N (U). KANE: Coral Pink Sand Dunes (BAD), Kanab 10 mi N (159) and 20 mi N (C42), Zion Nat Park (159). MIL- LARD; Black Rock 8 mi N (KU). SALT LAKE: Alta, Big Cottonwood Cyn (RG), Brighton, Butterfield Cyn (U), City Crk Cyn, Lake Blanche (RG), Mt Olympus, Parleys Cyn, Red Butte Cyn (U), Salt Lake City (KU). SAN JUAN: Abajo Mts (G58), Blanding 15.6 mi N (A), Bluff (U), Mexican Hat 13 mi S (A), Nat Bridges Nat Mon (Wi), Red Mesa (BAD), White Cyn (Wi). SANPETE: Fairview Cyn (KU). SUMMIT: Kamas 14.7 mi E (A), Woodland (BAD). TOOELE: S Willow Cyn (U). UIN- TAH: Bonanza and 3 mi S (KU), Gusher, Jensen (U), Dry Fk rd 22.8 mi N U121 (A). UTAH: American Fk Cyn (C42), Diamond Fk Cyn (KU), Provo Cyn (U), Santaquin Cyn 3.6 and 7.1 mi up, Tibbie Fk Cyn, Tibbie Fk Lake 0.5 mi W (A), Wanrhodes Cyn (KU). WASATCH: Cas- cade Spngs 3 mi N, Francis 14.4 and 18.5 mi E (A), Fruitland 4 mi W (U), Hanna 14.3 mi W (A), Heber (Wi), Midway, Soapstone Cyn (U), Soldier Summit (US). WASHINGTON: Grafton, Kolob, Rockville, Santa Clara Crk, Veyo, Virgin River E Fk (159), Zion Nat Park (WU). WAYNE: Elkhorn Ranger Sta, Pleasant Crk (BAD). WE- BER: Beaver Crk (KU), Ogden (Wi). Ants that run to niger in the key should be considered as ''niger complex," for some specimens whose records are included here are lighter in color and/ or larger than the "typical" niger, and may represent a different species. Smith (1979:1436) lists this species from the western United States, including Utah, Colorado, Arizona, and Idaho, nesting under stones and rotten wood in forests or open sit- uations. Gregg (1963:459) lists it from Colo- rado between 3500 and 12,400 ft under rocks and logs in a variety of habitats, pre- dominantly conifers. La Rivers (1968:8) lists it from Nevada, where Wheeler and Wheeler (1978:393) found it between 7600 and 9000 ft. They found it frequently in soil craters, and, in North Dakota, also common under rocks (1963). Cole (1942:374) states that in Utah its habitat is under stones and logs at higher elevations to 10,000 ft. Grundmann (1958:166) indicates that in Utah it nests un- der stones and rotting wood alongside streams between 5000 and 10,000 ft. Ingham (1959, 1963) found it in southern Utah under stones and logs, and in flat craters in open areas in a variety of vegetative types. There were 188 ants in 13 collections taken from under rocks. Lasius subumbratus was under the same rock in two collections, L. sitiens in one, and L. alienus and Formica obscuriventris in one. Six ants in one collec- tion were taken singly in an open area. Eggs were found under one rock in early July, lar- vae under one in late June, pupae under an- other in late June, and winged forms under one rock in late July. Eight collections were in aspen: two in association with grass, sage- brush, and snowberry; two maple, oak, and fir; one sagebrush; and one fir. Three collec- tions were in sagebrush: one in association with snowberry and oak, and one with herbs and conifers. One collection was in a grassy meadow with oak and conifers, and one in 482 Great Basin Naturalist Vol. 42, No. 4 ephedra and blackbrush. In 79 recorded Utah habitats it occurred 41 times in montane areas. In 51 elevational records between 3500 and 10,000 ft it was somewhat equally dis- tributed between 4000 and 8000, once over 10,000. Beck et al. (1967:70) found it feeding on dead rodents in seven instances in Utah. When the protective rock is removed, these ants run rapidly into burrows and to the opposite side of the rock. Larvae that are present may be abandoned with no attempt on the part of the workers to carry them into deeper burrows. Sometimes those on the ground stay and work furiously to move the eggs and pupae into the burrows. Under one rock three ApJiaenogaster siibterranea were present that had antennae and legs missing. These likely were captured by niger. There were 124 ants in six collections taken from under rocks. In one case these ants were under the same rock as Aphaeno- gaster siibterranea, and once with L. sub- iimbratus. In this latter case the tunnels of the two species of Lasius were distinctly apart under separate areas of the boulder. Two ants in one collection were taken singly away from the colony. Three collections were in aspen, one in association with con- ifers. One collection was in ephedra and blackbrush, one herbs, one a grassy meadow, and one firs. In 34 recorded Utah habitats it was taken 19 times in montane forest. In 15 recorded elevations between 4276 and 8750 ft it was taken most frequently under 6000. Beck et al. (1967:71) found it feeding on dead rodents in eight instances in Utah. Lasius pallitarsis (Provancher) Formica pallitarsis Provancher, 1881, Nat. Canad. 12:355. Lasius niger var. sitkaensis: Rees and Grundmann 1940:7; Cole 1942:374. Lasius sitkaensis: Beck et al. 1967:71. Lcptothorax sitkaensis: Knowlton 1975:6. Records (Map 16): BOX ELDER: SnowviUe (C42). CACHE: Bear River Range, Blacksmith Fk Cyn, Frank- lin Basin, Logan, Logan Cyn (KU). CARBON: Scofield (BAD). DAGGETT: Red Crk (BAD). DUCHESNE: Duchesne 3 mi E (WU). GARFIELD: Brvce Cyn Nat Park (WU). KANE: Kanab (C42). RICH: Meadowville (KU). SALT LAKE: Big Cottonwood Cyn, Holladay, Mill Crk Cyn (C42), Red Butte Cvn (U), Salt Lake City (C42). SAN JUAN: La Sal (RG), Mexican Hat 13 mi S (A). SANPETE: Bluebell Flats (KU), Ephraim 8.8 mi E (A), Mt Pleasant (BAD), Orangeville 27.4 mi W (A). SE- VIER: Fish Lake (BY), Koosharem (BAD). UINTAH: Bo- nanza (KU), Dry Fk rd 22.8 mi N U121 (A), Gusher (C42), Red Cloud Loop rd 9.2 mi W U44 (A). UTAH: Aspen Grove (BAD), Santaquin Cvn 3.6 mi up (A). WASATCH: Wallsburg (BAD). WASHINGTON: Zion Nat Park (C42). WAYNE: Pleasant Crk (BAD). Smith (1979:1437) lists this species from eastern to western United States, including Arizona and Nevada, nesting under stones or logs in forested areas. Gregg (1963:463) lists it from Colorado between 4600 and 12,200 ft under rocks and logs in a variety of habitats, predominantly conifers. Wheeler and Wheel- er (1978:393) found it between 6000 and 9700 ft in Nevada, and frequently under rocks, also common in and under wood and in soil in craters in North Dakota (1963). Cole (1942:374) indicates its habitat in Utah as under stones. Lasius sitiens Wilson L. sitiens Wilson, 1955, Bull. Mus. Comp. Zool. 113:108. Records (Map 17): CACHE: Logan (KU). JUAB: Fish Spngs Ranch (U), Topaz Mt (KU). UTAH: Diamond Fk Cyn (KU), Tibbie Fk Cyn (A), Wanrhodes Cyn (KU). Smith (1979:1437) lists this species from the western United States, including Colo- rado, Arizona, and Nevada, nesting under stones in dry open areas between 7000 and 8000 ft. Cole (1966:20) found its nests under stones in pinyon-juniper in southern Nevada. Thirty ants in one collection were taken from under a rock in an association of oak, maple, aspen, and fir in Utah. Lasius subumbratus Viereck L. umbratus subumbratus Viereck, 1903, Trans. Amer. Ent. Soc. 29:73; Rees and Gmndmann 1940:7; Cole 1942:375. L. subumbratus: Creighton 1950:424. Records (Map 16): DUCHESNE: Mirror Lake (Wi). GARFIELD: Bryce Cyn Nat Park (Wi). GRAND: Warner Ranger Sta (Wi). KANE: Cedar City 24.3 mi E (A), Long Valley Mts (Wi). SALT LAKE: Big Cotton- wood Cyn, Brighton (U), Lake Blanche (Wi). SAN JUAN: Blanding 16.5 mi N (A), Blue Mts (Wi) (? = Abajo Mts). SANPETE: Ephraim 8.8 mi E (A). SUM- MIT: Kamas 4.6 and 9.2 mi E, Mirror Lake 11.6 mi N (A), Shingle Crk (Wi), UINTAH: Red Cloud Loop rd 9.2 mi W U44 (A). UTAH: Orem (A), Provo (U), Tibbie Fk Lake (A), Timpanogos Pk (Wi). WASATCH: Cascade Spngs 3 mi N (A). WEBER: Woodruff 29.1 mi W (A). Smith (1979:1439) lists this species from eastern to western United States, including Arizona and Nevada, nesting under stones or December 1982 Allred: Ants of Utah 483 logs in meadows and forests. It is a social par- asite of L. pallitarsis. Gregg (1963:477) lists it from Colorado between 5160 and 9224 ft un- der rocks and logs in a variety of habitats. Cole (1942:375) indicates it as uncommon in Utah, nesting under stones. There were 294 ants in nine collections taken from mider rocks. In two instances ants of L. niger were imder the same rocks, and once those of L. pallitarsiis were present. Ten ants in one collection were found imder a log, and 20 ants in one collection singly in a garden. Six collections were taken in co- nifers: two in association with grass, herbs and aspen; two sagebrush and aspen; one a grassy meadow; and one aspen. Two collec- tions were taken in grass; one sagebrush; one sagebrush, snowberry, and oak; and one a garden. In 22 recorded Utah habitats it was taken 19 times in montane forest. Six record- ed elevations were between 5000 and 10,050 ft. Lasiiis umbratus (Nylander) Fonnka iimbrata Nylander, 1846, Acta. Soc. Sci. Fenn. 2:1048. L. umbratus mixtus var. aphidicohr. Rees and Gnind- mann 1940:7; Cole 1942:375. L. umbratus aphidicola: Ingham 1959:74. L. umbratus: Smith 1979:1439. Records (Map 16): BEAVER: Beaver 5.5 mi E (U). CACHE: Logan Cvn (C42). IRON: Cedar Citv 19 mi E (A). RICH: Woodruff (U). SALT LAKE: Big Cotton- wood Cyn (U), Little Willow Crk Cyn (RG). SAN JUAN: Kigalia Ranger Sta (Wi). SANPETE: Pine Plan- tation (KU). SUMMIT: Henefer (C42). UINTAH: Jensen (U). UTAH: Jordan Narrows (C42), Santaquin Cyn 7.1 mi up (A). WASHINGTON: Kolob (159). WEBER: Og- den Cyn (C42). Smith (1979:1439) lists this species from eastern to western United States, including Utah, Arizona, and Idaho, nesting under stones or logs. It is associated with L. alienus and L. niger. Gregg (1963:478) lists it from Colorado between 5254 and 9500 ft under rocks predominantly in conifer habitats. Wheeler and Wheeler (1963) found it fre- quently under rocks and in earthen mounds in North Dakota. Cole (1942:375) indicates its habitat in Utah as under stones. Ingham (1959) found it in logs in oak and aspen in southern Utah. Forty ants in one collection were found under a log, and 40 in another collection un- der a rock. Two collections were taken in aspen: one in association with pine and one in a grassy meadow. In 14 recorded Utah habitats it was taken 10 times in montane forest. Six recorded elevations were between 4494 and 8402 ft. Lasius vestitus Wheeler L. umbratus vestitus Wheeler, 1910, Psvche 17:238. Record: SAN JUAN; La Sal Mts (U).' Smith (1979:1440) lists this species from the western United States, including Idaho. Wheeler and Wheeler (1978:393) found it be- tween 7600 and 8100 ft in Nevada. Leptothorax ambiguus Emery L. cunispinosus ambiguus Emery, 1895, Zool. Jahrb. Syst. 8:317. Records: BOX ELDER: Locomotive Spngs (KU). UTAH: Spanish Fk Cyn (KU). Smith (1979:1392) lists this species from midwestern United States; no intermountain state is listed. It nests in soil or hollow grass stems in woodlands and grasslands. Its occur- rence in Utah is doubtful. Leptothorax andrei Emery L. andrei Emerv, 1895, Zool. Jahrb. Syst. 8:318. Record: TOOELE: Granite Mt (U). Smith (1979:1392) lists this species from western United States, including Arizona and Nevada, nesting under stones. Cole (1966:17) found it in southern Nevada in pinyon-juni- per, where it probably nests under stones. Allred and Cole (1971:239) found it in juni- per in Idaho. Leptothorax crassipilis Wheeler L. acervorum crassipilis Wheeler, 1917, Proc. .\mer. Acad. Arts. Sci. Bo.ston 52:513. L. crassipilis: Creighton 1950:278; Smith 1979:1396. Record: CARBON; Scofield 4 mi S (A). Smith (1979:1396) lists this species from western United States, including Utah, Colo- rado, Arizona, and Wyoming, nesting under rocks and logs. Gregg (1963:402) lists it be- tween 5700 and 9100 ft under rocks and logs in conifers, oak, and manzanita habitats in Colorado. La Rivers (1968:6) lists it from Ne- vada, where Wheeler and Wheeler (1978:391) found it at 8100 ft. Three ants in one collection in Utah were taken from un- der an aspen log. 484 Great Basin Naturalist Vol. 42, No. 4 Leptothorax furunculus Wheeler L. funinculus Wheeler, 1909. J. New York Ent. Soc. 17:82; Knowlton I975;5. Records: BOX ELDER: Cedar Crk (City), Curlew Jet, Hansel Mts, Snowville (K75), Wellsville Mts (KU). Smith (1979:1393) lists this species from Colorado and Wyoming. Gregg (1963:383) lists it from Colorado between 6970 and 7500 ft in pinyon-juniper areas. Knowlton (1975:5) found it associated with sagebrush in north- ern Utah. Leptothorax hirticomis Emery L. hirticornis Emery, 1895, Zool. Jahrb. Svst. 8:317; Cole 1942:370; Smith 1979:1397. Record; SALT LAKE: locality unknown (C42). Smith (1979:1397) lists this species from midwest and western United States, including Utah and Colorado, nesting with Formica ob- scuripes in large mounds of detritus in open areas. Gregg (1963:405) lists it between 5354 and 7000 ft in Colorado. Leptothorax muscorum (Nylander) Myrmica muscorum Nvlander, 1846, Acta Soc. Sci. Fenn. 2:1054. L. acervoTum canadensis: Rees and Gnmdmann 1940:6; Cole 1942:369; Hayward 1945:120. L. acervorum canadensis var. yankee: Rees and Gnmd- mann 1940:6; Cole 1942:369. L. canadensis: Ingham 1959:60. L. muscorum: Beck et al. 1967:71. Records (Map 17): BOX ELDER: Box Elder Cyn (US). CACHE: Blacksmith Fk Cyn, W Hodges Cyn (KU), Lo- gan (C42), Logan Cyn Summit, Mendon Cold Spngs (KU), Millville (US), tony Grove, Wellsville (KU). GAR- FIELD: Osiris (U). IRON: Cedar Breaks Nat Mon (159). JUAB: Red Crk Spng (KU). SALT LAKE: Big Cotton- wood Cyn S Fk (C42), Brighton, Butterfield'Cyn (U), Little Cottonwood Cyn (C42). SANPETE: Manti (KU). UINTAH: Paradise Park (U), Whiterocks Cyn (KU). UTAH: American Fk Cyn (U), Aspen Grove (BY), Emer- ald Lake (BAD). Smith (1979:1397) lists this species from eastern to western United States, including Arizona, nesting imder rocks, logs, or bark of fallen trees in woodlands. Gregg (1963) lists it from Colorado between 5354 and 12,500 ft imder rocks and logs in a variety of habitats, predominantly in conifers. La Rivers (1968:6) lists it from Nevada, where Wheeler and Wheeler (1978:391) found it between 6400 and 11,000 ft. Cole (1942:369) indicates its habitat in Utah as imder decaying wood and in standing dead trees. Ingham (1959) found it under logs in fir, spruce, and bristlecone pine in southern Utah. In 22 recorded Utah habitats it was found 15 times in montane forest. In 16 recorded elevations between 4495 and 10,500 ft it was taken most frequently above 7000. Beck et al. (1967:71) found it feeding on dead rodents in one instance in Utah. Leptothorax nevadensis Wheeler L. nevadensis Wheeler, 1903, Proc. Acad. Nat. Sci. Phila. 55:224; Rees and Gnmdmann 1940:6; Cole 1942:370; Knowlton 1970:210, 1975:5. Records (Map 17); BEAVER: Beaver 5.5 mi E (U). BOX ELDER: Cedar Crk (City) (K70), Curlew Valley (US), Hansel Mts, Kelton Pass, Snowville (K70). CACHE: Blacksmith Fk Cyn (RG), Green Cvn (US). RICH: Randolph 8 mi SW (KU). SANPETE: Ephraim Cvn (KU). Creighton (1950) and Smith (1979:1393) list four forms of this species from the west- ern United States, including Nevada, nesting in soil usvially under stones. The subspecies that occurs in Utah is likely rudis or neva- densis. These can be separated by the sculp- ture on the dorsum of the thorax. On neva- densis the thorax is densely and evenly punctate, whereas on rudis the punctures are interrupted by prominent rugae on the epinotum and mesonotum. Cole (1966:17) found its nests in southern Nevada in pinyon- juniper and under stones in Utah (1942:370). Wheeler and Wheeler (1978:392) found it be- tween 6100 and 10,000 ft in Nevada. Knowl- ton (1975:5) found it associated with rabbit- brush and grass in northern Utah. In 10 recorded Utah habitats it was taken four times in montane forest. One recorded eleva- tion is 4544 ft. Leptothorax nitens Emery L. nitens Emery, 1895, Zool. Jahrb. Syst. 8:318; Rees and Grundmann 1940:6; Cole 1942:370; Creighton 1950:265; Grundmann 1958:164; Knowlton 1970:211, 1975:6. Records (Map 18): BOX ELDER: Box Elder Cyn (KU), Cedar Crk (City), Hansel Mts (K75), Kelton, Kel- ton Pass, Snowville (k70) and 13 mi SW, Wellsville Mts (KU). CACHE: Providence Cyn (C42), Spring Hollow (US). GARFIELD: Boulder Mt (G58). GRAND: Moab 10 mi SE (KU). MILLARD: Black Rock (US). SAN JUAN: Abajo Mts (G58). UTAH: American Fk Cyn (RG). December 1982 Allred: Ants of Utah 485 Smith (1979:1394) lists this species from western United States, inchiding Colorado and Wyoming, nesting under rocks and in duff. Hunt and Snelling (1975:22) list it from Arizona. Gregg (1963:384) lists it between 6000 and 8000 ft imder rocks in conifers, pin- yon-juniper, and oak habitats in Colorado. La Rivers (1968:6) lists it from Nevada, where Wheeler and Wheeler (1978:392) found it at 6300 and 6400 ft. Cole (1942:370) found it under stones in Utah. Gnmdmann (1958:164) indicates that it nests under stones in desert conditions and transition zones in shrubs in Utah. Knowlton (1975:6) found it associated with horsebnish in northern Utah. In 15 re- corded Utah habitats between 4225 and 7000 ft, it was found 7 times in montane areas. Leptothorax rugatulus Wheeler L. rugatulus brunnescens Wheeler, 1917, Proc. Amer. Acad. Arts, Sci. 52:510; Creighton 1950:269; Smith 1979:1394. L. rugatulus: Cole 1942:369; Creighton 1950:268; Grundmann 1958:164; Knowlton 1975:6. Records (Map 18): BOX ELDER: Box Elder Cvn (KU), Cedar Hill (K75), Park Valley (City) (C42), Snow- ville (K75) and 13 mi SW (KU), Wildcat Hills (K75). CACHE: Blacksmith Fk Cyn (US), Green Cyn (KU), Hyde Park (US), Logan Cyn (KU), Richmond (US). MIL- LARD: White Valley (C'42). SALT LAKE: Red Butte Cvn (U). SAN JUAN: Mexican Hat (G58). SANPETE: Majors Flats (KU). TOOELE: Clover, Delle, Fisher Pass (C42). UINTAH: Bonanza 3 mi S (KU). WEBER: Uintah (US). Creighton (1950) and Smith (1979:1394) list two races of this species from the western United States, including Utah and Colorado, where it nests under rocks or wood. The Utah race is likely the subspecies brunnescens, which may be separated from rugatulus by the thoracic rugae that are well developed on rugatulus, but feeble, often replaced by pimctures, on brunnescens. Himt and Snell- ing (1975:23) list it from Arizona. Gregg (1963:387) lists it from Colorado between 5354 and 8700 ft under rocks and logs in a variety of habitats, predominantly conifers. La Rivers (1968:6) lists it from Nevada, where Wheeler and Wheeler (1978:392) found it between 6000 and 10,000 ft. They foimd it under rocks and other objects in North Dakota (1963). Cole (1942:369) in- dicates that it nests under stones in Utah. In Utah it occurs in the transition zone (Grund- mann 1958:164). Knowlton (1975:6) found it associated with sagebrush and snowberry in northern Utah. In 20 recorded Utah localities it occurred 6 times in montane areas between elevations of 4270 and 6300 ft, mostly at lower elevations. Leptothorax silvestrii (Santschi) Tetramoriunt silvestrii Santschi, 1909, Soc. Ent. Ital., Bol. 41:6. Record: BOX ELDER: Cedar Hill (KU). Smith (1979:1395) lists this species from southern Arizona nesting in oak above 3500 ft. Its occurrence in Utah is questionable. Leptothorax tricarinatus Emery L. tricarinatus Emerv, 1895, Arb. Zool. Jahrb. System 8:318; Knowlton 1970:211, 1975:6. L. tricarinatus tricarinatus: Smith 1952:100; Smith 1979:1395. L. tricarinatus neoinexicanus: Smith 1952:101; Grund- mann 1958:164; Ingham 1959:60; Smith 1979:1395. Records (Map 18): BOX ELDER: Kelton (K70). DUCHESNE: Avintaquin Cmpgnd (A). JUAB: Mt Nebo (US), Nephi 20 mi SW (Sm52). MILLARD: White Val- ley (Sm52). SANPETE: Orangeville 19.5 mi W (A). SUMMIT: Mirror Lake 17.3 mi N (A). UINTAH: Red Cloud Loop rd 19 mi W U44 (A). WASATCH: Daniels Pass (US). WASHINGTON: Kolob (159). Creighton (1950) and Smith (1979:1395) list two races of this species from western United States, including Utah, Colorado, Ari- zona, and Wyoming, nesting in soil and un- der rocks in open grassy areas. Both races are known for Utah. The subspecies neo- mexicanus may be separated from tricari- natus by the length of the epinotal spines, which are longer on neomexicanus, and by the opaque thorax of neomexicanus versus the shining surface of tricarinatus. Gregg (1963) lists it between 4600 and 7800 ft un- der rocks in a variety of habitats, pre- dominantly grass areas in Colorado. Gnmd- mann (1958:164) lists it as a plateau form between 6000 and 8000 ft in Utah. Ingham (1959) found it in clumps of dry grass associ- ated with sagebrush, oak, and a variety of herbs in southern Utah. There were 120 ants in three collections takeji from under rocks, and 50 in one collec- tion under a log. Immatures were found un- der one rock in early July. Two collections were taken in aspen: one in association with fir and one with sagebrush and conifers. One 486 Great Basin Naturalist Vol. 42, No. 4 collection was taken in grass and one in pine. It was found in 7 montane areas in 10 local- ities where the habitat was recorded, at ele- vations from 4225 to 8000 ft. Liometopum occidentale Wheeler L. apiculatum luctuosum Wheeler, 1905, Bull. Amer. Mus. Nat. Hist. 21:.325; Rees and Griindmann 1940:6; Cole 1942:371. L. occidentale hictiiostan: Creighton 19.50:339; Grund- niann 1958:16.5; Ingham 19.59:60. L. tricarinatiis: Knowlton 1970:211, 1975:6. L. luctuosum: Allred and Cole 1979:99. Records (Map 18): CARBON: Wellington 10 mi NE (A). DUCHESNE: Duchesne 9 mi W (WU), Myton (RG). EMERY: Greenriver, Gunnison Butte (RG). GRAND: Dewey (U), Moab (RG). KANE: Glen Cyn City (AC), Kanab 12.5 mi N (A), Mt Carmel Jet, Zion Nat Park (159). PIUTE: Marysvale 4.1 mi S (A). SAN JUAN: Blanding (G58), Hatch Wash nr La Sal (RG), Monticello 17 mi E (U). WASHINGTON: Rockville, Zion Nat Park (159). WAYNE: Capitol Reef Nat Park (U). Creighton (1950) and Smith (1979:1417) indicate two forms of this western United States species, including Colorado, Arizona, and Wyoming. The Utah specimens probably belong to the subspecies luctuosum rather than occidentale. Gregg (1963:443) lists this species between 4800 and 7550 ft under rocks and logs in conifers and pinyon-juniper in Colorado. Wheeler and Wheeler (1978:392) found it between 5200 and 8100 ft in Nevada. Cole (1966:18) found a nest in southern Nevada in detritus at the base of a juniper; it also nests under stones in Utah, for the most part at elevations above 4000 ft (1942:371). Grundmann (1958:165) indicates this as a mountain species that nests between 4000 and 7000 ft in Utah. Ingham (1959) found it in southern Utah under rocks and bark of living trees in a variety of habitats. Allred and Cole (1979:99) found it in sage- brush-grass and juniper-pinyon associations in southern Utah. Sixty ants in two collections were taken from small crater mounds, 29 in one collec- tion from under a rock, and 25 in one collec- tion from duff under a pinyon tree. Three collections were taken in sagebrush, one in association with juniper and pinyon. None of 18 recorded Utah localities were in montane areas. In 15 recorded elevations between 3250 and 7300 ft nine are between 4000 and 6000. Manica hunteri (Wheeler) Mynnica hunteri Wheeler, 1914, Psyche 21:119. Manica hunteri: Wheeler and Wheeler 1970:1:35; Smith 1979:1:352. Record: CACHE: Benson (W70). Smith (1979:1352) lists this species from the western United States, including Utah, Nevada, Idaho, and Wyoming, nesting in open areas of coniferous forests. Wheeler and Wheeler (1970) state that its habitat is under stones in craters in coniferous forest, specifi- cally lodgepole pine, although craters are rare. They found it between 2200 and 9600 ft in Nevada (1978:391). Manica niutica (Emery) Mijrmica mutica Emery, 1895, Zool. Jahrb. Syst. 8:311; Rees and Grundmann 1940:5; Cole 1942:368. Monica mutica: Ingham 1959:.38; Wheeler and Wheeler 1970:159; Knowlton 1970:211, 1975:6. Records (Map 19): BOX ELDER: Blue Crk (City) (C42), Cedar Hill (K70), Corinne (C42), Kelton, Penrose 7 mi W (KU), Tremonton (C42). CACHE: Benson (W70), Hyrum (C42), Lewiston (US). DAVIS: NE Ante- lope Island (W70), Lavton (W70), Woods Cro.ss (RG). GARFIELD: Osiris, Panguitch (W70). IRON: Cedar Breaks Nat Mon (W70), Cedar City (RAU), Coal Crk Cyn (159). JUAB: Lexington (US). KANE: Long Valley (W70). MILLARD: Delta (BY), Fillmore (US). PIUTE: Junction (W70). SALT LAKE: Big Cottonwood Cyn (U), Great Salt Lake S end, Midvale, Murray (W70), Salt Lake City (RG). SANPETE: Chester, Fayette (C42). TOOELE: losepa. Lake Point, Tooele (C42). UTAH: As- pen Grove (BY), Lehi (W70), Proyo (U), Salem, Spring- ville (C42), Thistle 14.6 mi E (A), E Utah Lake (BY). WEBER: Ogden, Plain City (C42). Smith (1979:1352) lists this species from midwest and western United States, including Colorado, nesting in a wide variety of habi- tats, where it is associated with Formicoxenus chamberlini. Gregg (1963:316) lists it from Colorado between 4500 and 8600 ft under rocks and wood in conifers, pinyon-juniper, cottonwood-willow, grass, sagebrush, and greasewood habitats. La Rivers (1968:2) lists it from Nevada. Wheeler and Wheeler (1970) list it from Colorado, Arizona, Nevada, Idaho, Wyoming, and Utah, where it often nests in small craterlike mounds in a variety of habitats including coniferous forest, pin- yon-juniper, and grasslands between 1100 and 8600 ft. They found it frequently in soil craters in North Dakota (1963). Cole (1942:368) indicates that in Utah it nests un- der stones and in ditch banks, sometimes con- structing small craters. Ingham (1959, 1963) December 1982 Allred: Ants of Utah 487 found it under rocks in juniper and alkali flats in southern Utah. Twenty ants in one collection in Utah were taken from imder a rock in an associ- ation of grass, clover, Russian thistle, and sagebrush. In 41 recorded habitats it was taken only six times in montane areas. In 35 recorded elevations between 4225 and 10,399 ft it was taken most frequently under 5000, once over 8000. Monomorium minimum (Buckley) Myrmica minima Buckley, 1867, Proc. Ent. Soc. Phila. 6:338. Monotnorium minimum: Rees and Gmndmann 1940:3; Cole 1942:361; Grundmann 1958:163; Ingham 1959:57; Beck et al. 1967:71; Knowlton 1975:6; Allred and Cole 1979:98. Records (Map 19): BEAVER: Beaver 5.5 mi E (U). BOX ELDER: Hansel Mts (K75), Locomotive Spngs (KU). CACHE: Ant Vallev, Green Cyn, Leeds Cyn (KU). EMERY: Huntington (BAD). GARFIELD: Boulder (U). GRAND: Moab 15 mi S (KU). IRON: Highway US91 (159). JUAB: Callao (BAD). KANE: Glen Cyn City (AC), W Glen Cyn Res (G58), Grosvenor Arch' (US), Kanab (G58) and 20 mi N (C42), Long Valley Jet 2 mi W, Mt Carmel Jet (159). MILLARD: Swasey Spngs (RG). MOR- GAN: Morgan (KU). SALT LAKE: Big Cottonwood Cvn (U), Little Willow Cvn (C42), Rose Cyn (U), Salt Lake City (RG). SAN JUAN: Blanding (RG), Mexican Hat (G58). SANPETE: Manti (KU). SEVIER: Sevier (KU). SUMMIT: Echo (BAD). TOOELE: Clover, Fisher Pass, Little Valley Ranger Sta, Orrs Ranch (C42). UTAH: .\merican Fk (RG), Aspen Grove (BY), Diamond Fk Cyn, Spanish Fk Cyn (KU). WASATCH: Provo River N Fk (U). WASHINGTON: Highway 18 (159), Pinto 3 mi S (US) and 3 mi N (KU), Zion Nat Park (159). WEBER: Hooper (C42). Smith (1979:1382) lists this species from Canada to Florida and western United States, including Colorado, with nests in exposed soil, imder objects, or in rotting wood. Hunt and Snelling (1975:21) list it from Arizona. Gregg (1963:368) Usts it between 3500 and 8500 ft under rocks predominantly in pinyon- juniper areas in Colorado. Cole (1966:16) found it in southern Nevada under stones in blackbrush and other desert shrub types, rarely in pinyon-jimiper. He states that in Utah it nests under stones, in crater mounds, vmder bark, or in logs (1942:361). Allred and Cole (1979:98, 1971:239) found it in southern Utah and Idaho in a variety of desert shnib types. Wheeler and Wheeler (1963) found it commonly in soil craters in North Dakota. Grundmann (1958:163) found it abundant in transition zones and canyon sides under stones in Utah. Ingham (1959, 1963) found it under stones or in .small crater mounds in open areas associated with sagebrush, juni- per, thistlepoppy, galletagrass, little rabbit- brush, winterfat, and greasewood in southern Utah. Knowlton (1975:6) found it associated with rabbitbrush in northern Utah. In 43 recorded Utah habitats only 10 were in montane areas. Twenty-seven elevational records between 3250 and 8000 ft show it most common (24 times) between 4000 and 7000. Beck et al. (1967:71) found it feeding on dead rodents in three instances in Utah. Myrmecocystus flaviceps Wheeler M. ijiima var. flaviceps Wheeler, 1912, Psyche 19:174. M. flaviceps: Snelling 1976:85; Smith 1979:1446. Records (Map 19): JUAB: Callao 5 mi E (U). MIL- LARD: Black Rock (US) and 5 to 8 mi N, Deseret 30 to 32 mi S (S76). WASHINGTON: Harrisburg (S76), Zion Nat Park (WU). Smith (1979:1446) lists this species from western United States, including Utah, Ari- zona, and Nevada. Snelling (1976:85) in- dicates that its nests are craterlike in juniper, sagebrush, saltbush-greasewood, creosote bush, creosote bush-bur sage, and palo verde- cactus. Myrmecocystus ham.mettensis Cole M. hammettei^sis Cole, 19,38, Amer. Midi. Natl. 19:678. Record: BOX ELDER: Park Valley (City) 31 mi SW (KU). Smith (1979:1446) lists this species from western United States, including Nevada and Idaho. Snelling (1976:102) indicates its habi- tat as saltbush-greasewood and sagebrush- grass in craterlike nests. Myrmecocystus kennedyi Cole M. melliger semirufus var. kennedi/i Cole, 1936, Ent. News 47:119. M. melliger semirufus: Cole 1942:386. M. kennedyi: Creighton 1950:449; Grundmann 1958:167; Snelling 1976:65; Smith 1979:1446. Records: BOX ELDER: Lucin (C42). GRAND: Moab (C42). JUAB: Silver City (US). WASHINGTON: St George 5 mi S (S76). Smith (1979:1446) lists this species from western United States, including Utah, Ari- zona, Nevada, and Idaho. Snelling (1976:66) lists its habitats as sagebrush, creosote bush- bur sage, and palo verde-cactus, where nests 488 Great Basin Naturalist Vol. 42, No. 4 occur as craters in bare areas away from veg- etation. Cole (1942:386) indicates its habitat as craterlike nests in open areas in Utah. None of four recorded Utah habitats were in montane areas. Four recorded elevations were between 4000 and 6100 ft. Myrmecocystus mendax Wheeler M. meUiger mendax Wheeler, 1908, Bull. Anier. Mus. Nat. Hist. 24:351; Rees and Griindmann 1940:11; Cole 1942:385. M. melliger. Allred and Cole 1979:98. Records: CACHE: Wellsville (RG). EMERY: Green- river (US). KANE: Glen Cyn City (AC). MILLARD: Swasey Spngs (C42). SAN JUAN: Dead Horse Pt State Park (US). Smith (1979:1447) lists this species from western United States, including Colorado, Arizona, and Nevada. Gregg (1963:645) lists it between 3600 and 6600 ft under rocks and in crater nests in pinyon-juniper and grass habitats in Colorado. Snelling (1976:40) in- dicates its habitats as pinyon-oak, mesquite- acacia, pinyon-juniper, oak-juniper, and shrubs between 3600 and 6600 ft. Cole (1942:386) indicates that in Utah its nests are in unprotected soil of open areas. Allred and Cole (1979:98) found it in a variety of desert shrub types in southern Utah with no appar- ent plant preference. None of the recorded habitats in Utah were from montane areas, and four recorded elevations were between 3250 and 6000 ft. Myrmecocystus mexicanus Waesmael M. mexicanus Wesmael, 1838, Bull. Acad. Sci. Belg. 5:770; Garrett 1910:342; Snelling 1976:122; All- red and Cole 1979:98; Smith 1979:1447. M. mexicanus hortideorum: Rees and Grundmann 1940:11; Cole 1942:386; Creighton 1950:446; Gnindmann 1958:166; Ingham 1959:75; Beck et al. 1967:71. Records (Map 20): EMERY: Green River, Gunnison Butte (RG). GARFIELD: Boulder, Henry Mts (S76), Shootering Cyn (US). GRAND: Arches Nat Park (S76), Moab (G58), Thomp.son (C42). IRON: Shirts Cyn (RAU). KANE: Glen Cyn City (AC). MILLARD: Black Rock (US) and 5 mi N (KU), Deseret 32 mi S (S76). SALT LAKE: Garfield (S76). SAN JUAN: Aztec Cyn (U), Blanding (G58), Bluff (RG), Four Corners, Mexican Wa- ter, Montezuma Crk, Red Mesa (BAD). WASHING- TON: Hurricane 3 mi SW (159), Pine Valley (Citv) (BAD), Santa Clara (US), St George (BY). Smith (1979:1447) lists this as a western species of United States, including Utah, Col- orado, Arizona, and Nevada. Gregg (1963:648) lists it between 4500 and 6800 ft in large craters predominantly in pinyon- juniper and sagebrush habitats in Colorado. In southern Nevada Cole (1966) found it abundantly in hopsage-matrimony vine, Rus- sian thistle, blackbrush, and pinyon-juniper habitats. It was scarce in creosote bush areas. Nests in Utah are craterlike, generally on hills and ridges with sparse cover (Cole 1942:386). Grundmann (1958:166) indicates it as a desert species in Utah that occasionally may be found chewing on the ears of mice caught in traps. Ingham (1959, 1963) found it in southern Utah in a variety of shrub types, and Allred and Cole (1979:98, 1971:239) found it in southern Utah and Idaho in a vari- ety of desert shrub types. Snelling (1976:124) indicates that its nests are craterlike. In 31 recorded Utah habitats it was never found in a montane area. In 18 elevational records it was about equally distributed be- tween 2625 and 6675 ft, slightly more com- mon above 4000. Beck et al. (1967:71) found it feeding on dead rodents in 13 instances in Utah. Myrmecocystus mimicus Wheeler M. meUiger mimicus Wheeler, 1908, Amer. Mus. Nat. Hist., Bull. 24:353. M. mimicus: Ingham 1959:76; Allred and Cole 1979:99. Records: CACHE: Green Cyn (KU). WASHINGTON: Grafton, Harrisburg Jet 2 mi W, Harrisburg Jet toward Hurricane (159). Smith (1979:1447) lists this species from midwest to western United States, including Arizona. Cole (1966:22) found it commonly in creosote bush habitats in southern Nevada, as well as some other desert shrub types. The nests are holes in the ground or in craters in open areas. Ingham (1959, 1963) found it in southern Utah in craters in bur sage, rabbit- brush, cholla, creosote bush, mesquite, sage- brush, Joshua trees, sand sage, four-wing salt- bush, and shadscale. Allred and Cole (1979:99) found it in juniper-ephedra-grass, ephedra-grass, and blackbrush associations in southern Utah. Snelling (1976:58) lists pin- yon-juniper, oak-juniper, sagebrush, saltbush- greasewood, creosote bush-bur sage, and grassland at habitats below 4000 ft. Nests are craterlike, frequently concealed by a grass clump. Only one of four recorded Utah habi- tats was in a montane area. Five recorded December 1982 Allred: Ants of Utah 489 elevations were between 2700 and 5000 ft. Snelling (pers. comm.) believes that these Utah records are misidentifications. Myrtnecocysttis navajo Wheeler M. mexicanus ruivajo Wheeler, 1908, Bull. .\mer. Mus. Nat. Hist. 24:360; Cole 1942:386. M. navajo: Grundmann 1958:167; Snelling 1976:126; Smith 1979:1447. Records: BEAVER: Milford 26.7 mi W (A). JUAB: Trout Crk (City) 9 mi E (S76). MILLARD: Delta 60 mi W (S76), White Valley (C42). SAN JUAN: Bluff (G58). Smith (1979:1447) indicates this as a west- ern United States species, including Utah, Colorado, Arizona, and Nevada. Gregg (1963:651) lists it between 4100 and 4400 ft in craters in grass habitats in Colorado. Cole (1942:386) indicates that in Utah the nests are inconspicuous in open areas. Grundmann (1958:167) states that in Utah it inhabits ob- scure ground burrows, with the sand spread out in the form of a disc. Snelling (1976:128) indicates that nests are in the center of a flat disc or may be craterlike in saltbush-grease- wood. Fifty ants in one collection were taken from a crater moimd in shadscale in Utah. Myrrnecocystus pyramicus Smith M. pyramicus Smith, 1951, Great Basin Nat. 11:91; Beck et al. 1967:71. Record: SAN JUAN: Mexican Water (BAD). Smith (1979:1447) lists this species from western United States, including Nevada and Idaho. Snelling (1976:135) indicates its habi- tats as sagebrush and saltbush-greasewood, with craterlike nests between 2350 and 6700 ft. Beck et al. (1967:71) found it feeding on dead rodents in two instances in Utah. Myrrnecocystus romainei Cole M. melliger seniiritfus var. romainei Cole, 19.36, Ent. News 47:120. M. romainei: Snelling 1976:81; Smith 1979:1447. Records (Map 20): BOX ELDER: Park Valley (KU). EMERY: Greenriver (S76). GARFIELD: Dixie State Park (S76). IRON: Beryl (S76). KANE: Glen Cyn City (AC). MILLARD: Delta, Lynndyl (S76). SAN'JUAN: Bluff (S76). UINTAH: Dinosaur Nat Mon, Jen.sen (S76). WASHINGTON: Enterprise, Leeds (S76). Smith (1979:1447) lists this species from midwest to western United States, including Utah, Colorado, Arizona, and Nevada. Snell- ing (1976:81) designates its habitats as grass. pinyon-juniper, and creosote bush-tarbush in irregular crater nests. None of 12 recorded Utah habitats were in montane areas. In 11 recorded elevations between 2750 and 5600 ft, 9 were above 4000. Myrrnecocystus semirufus Emery M. melliger var. semirufus Emery, 1893, Zool. Jahrb. Syst. 7:667. M. semirufus: Grundmann 1958:167; Gregg 1963:653. Records: BOX ELDER: Park Valley (City) 31 mi SW (KU). DUCHESNE: Dinosaur Nat Mon (Gr63). Smith (1979:1448) lists this species as ex- treme western United States, but does not list an intermountain state. Snelling (pers. comm.) believes these Utah records to be misidentifications of M. kennedyi, M. men- dax, or M. romainei. Gregg (1963:653) lists semirufus from Colorado between 4800 and 5000 ft under rocks and in crater nests in pin- yon-juniper and cottonwood-willow habitats, and gives a record for Utah. La Rivers (1968:9) lists it from Nevada. Grundmann (1958:167) states that in Utah it lives in ground burrows surrounded by a sand crater. Snelling (1976:49) indicates its habitats as oakwoods, pinyon-juniper, sagebrush, creo- sote bush-bur sage, and creosote bush, with craterlike nests at elevations between 4000 and 5000 ft. Myrrnecocystus testaceus Emery M. melliger var. testaceus Emery, 1893, Zool. Jahrb. Syst. 7:667. M. testaceus: Beck et al. 1967:71; Snelling 1976:138; Smith 1979:1448. M. mojave: Beck et al. 1967:71. Records: BEAVER: Minersville (BAD). BOX ELDER: Lucin (BAD). CARBON: Price (BAD). EMERY: San Ra- fael River (BAD). JUAB: Callao (BAD). KANE: Adair- ville (BAD). MILLARD: Swasey Spngs (S76). SAN JUAN: Four Corners, Mexican Water, Montezuma Crk, Red Mesa (BAD). UINTAH: Duchesne 3 mi E (S76). WASHINGTON: Pine Valley (City) (BAD). WAYNE: Pleasant Crk (BAD). Smith (1979:1448) lists this species from western United States, including Utah, Ne- vada, and Idaho. Snelling (1976:139) in- dicates its habitats as sagebrush, pinyon-jimi- per, and chaparral between 1400 and 6900 ft in craterlike mounds. Four known specific habitats in Utah were in desert areas under 6000 ft. 490 Great Basin Naturalist Vol. 42, No. 4 Myrmica americana Weber M. sabuleti americana Weber, Lloydia 2:144; Rees and Gmndmann 1940:5; Cole 1942:368. M. americana: Creighton 1950:94; Grundmann 1958:162; Ingham 1959:36; Knowlton 1975:6; Smith 1979:1348. Records (Map 20): BOX ELDER: Cedar Hill (US), Hansel Mts, Kelton Pass (K75), Wildcat Hills (US). CACHE: Logan Cyn Summit (KU). GARFIELD: Bryce Cyn Nat Park (C42). KANE: Long Valley Jet 2 mi'w (159). RICH: Randolph 2.3 mi N, Sage Crk Jet 5.1 mi W (A). SALT LAKE: Big Cottonwood Cyn, Butterfield Cyn (U). SAN JUAN: Abajo Mts (G58). SANPETE: Orange- ville 19.5 mi W (A). SUMMIT: Kamas 21 and 26.4 mi E, Wyoming brdr on U150 (A). UINTAH: Bonanza 25 mi S (A). Smith (1979:1348) designates this as a com- mon grassland species more common east of the Rocky Mountains and rare in the west, but known from Utah, Colorado, and Ari- zona. Gregg (1963:310) lists it between 3500 and 7500 ft imder rocks and wood in conifers, pinyon-juniper, oak, mahogany, and grassy areas in Colorado. La Rivers (1968:2) lists it from Nevada. Wheeler and Wheeler (1978:389) found it at elevations as high as 10,400 ft in Nevada, and commonly under rocks in North Dakota (1963). Cole (1942:368) indicates that in Utah it nests un- der stones. Grundmann (1958:162) designates it as a mountain form from oak-jimiper to as- pen-fir, nesting under stones in Utah. Ingham (1959) foimd it under stones in southern Utah. Knowlton (1975:6) foimd it associated with rabbitbrush in northern Utah. Fifty-five ants in three collections in Utah were taken from under rocks, one under the same rock with Lasius crypticus. One collec- tion was in conifers, one grass, and one juni- per and piny on. In 17 localities 9 were in canyon-montane forest habitats. Five record- ed elevations were between 6500 and 7977 ft. Mymiica brevispinosa Wheeler M. rubra brevinodis var. brevispinosa Wheeler, 1907, Bull. Wis. Nat. Hist. Soc. 5:74. M. brevinodis discontintia: Beck et al. 1967:71. Records (Map 20): CARBON: Price (U), Scofield (BAD). EMERY: Carter Crk nr Green River (U). RICH: Laketown (BAD), Woodmff 13.8 and 18.4 mi W (A). SAN JUAN: Abajo Mts (U). TOOELE: Blue Lakes (30 mi S Wendover), Gold Hill (U). UINTAH: Ashley Crk nr Vernal, Paradise Park 11 mi S (U). WASHINGTON: Pine Valley (BAD). Creighton (1950) and Smith (1979:1348) list two races of this species from midwest to western United States, including Colorado, Idaho, and Wyoming, nesting near streams or permanent bodies of water. The Utah race is likely discontiniia, which may be distin- guished by its dark brown color in contrast to the orange yellow color of brevispinosa. Gregg (1963:299) lists this species between 4600 and 10,850 ft under rocks and wood in conifers, pinyon-juniper, and cottonwood- willow habitats in Colorado. Wheeler and Wheeler (1963) found it under rocks in North Dakota. Eighty ants in one collection were taken from under a rock, and one ant from under a log where Formica gnava was present. Two collections were in aspen, one in association with grass, sagebrush, and conifers. In 12 re- corded Utah habitats 4 were in montane for- est. Nine recorded elevations were between 5566 and 8000 ft. Beck et al. (1967:71) found it feeding on dead rodents in three instances in Utah. Myrmica emery ana Wheeler M. scabrinodis schenki var. einenjana Wheeler, 1917, Proc. Amer. Acad. Sci. Arts 52:504. M. scabrinodis sulcinodoides: Rees and Grundmann 1940:5. M. scabrinodis brevinodis: Rees and Grundmann 1940:5. M. schenecki eineryana: Cole 1942:368. M. scabrinodis mexicana: Cole 1942:368. M. emeryana tahoensis: Creighton 1950:99; Smith 1979:1349. M. emeryana emeryana: Grundmann 1958:161. M. emeryana: Ingham 1959:36. Records (Map 21): BEAVER: Beaver 5.5 mi E (U). CACHE: Franklin Basin, Leeds Cyn, Logan Cyn, Tony Grove Cyn (KU). GARFIELD: Aquarius'Plateau (G58), Boulder (U), Bryce Cyn Nat Park (RG), Escalante 20 mi E (U). KANE: Long Valley Jet 11 mi W (159). SALT LAKE: Big Cottonwood Cyn, Butterfield Cyn (U). SAN JUAN: Abajo Mts (U). SUMMIT: Soapstone Ranger Sta (U). UINTAH: Ashley Crk nr Vernal (U). UTAH: Aspen Grove (BY), Halls Fk rd 5.2 mi N Hobble Crk rd (A), Leki (? = Lehi) (C42), Mt Timpanogos (BY). Creighton (1950) and Smith (1979) desig- nate two races of this species occurring at high elevations in the mountains of western United States, including Utah, Arizona, Ne- vada, and Wyoming. The subspecies ta- hoensis may be separated from emeryana by the lamina of the antennal scape, which is small and diagonally transverse on emeryana. December 1982 Allred: Ants of Utah 491 but is a prominent median flange on ta- hoensis. Gregg (1963:312) lists this species from Colorado between 5800 and 9713 ft un- der rocks in conifers, oak, manzanita, and pinyon-jimiper areas. Cole (1966:3) states that in southern Nevada it is found under stones in pinyon-juniper. Wheeler and Wheeler (1978:389) found it at 9000 and 10,400 ft in Nevada and commonly in wood and under rocks in North Dakota (1963). Grundmann (1958:161) states that it is a mountain form in Utah, generally associated with aspen-fir above 5000 ft, and making nests imder stones. Ingham (1959) found it under stones in grass in southern Utah. One ant was collected under a rock in a meadow of grass, sedges, and herbs. Other ants under the same rock were Formica fiis- ca, F. gnava, and Mijmiica monticola. In 18 of 19 collections in Utah it was taken in can- yon-montane forest. The 12 recorded eleva- tions varied between 4562 and 8000 ft, 9 above 7000. Myrmica hamiihta Weber M. sahiileti hamulata Weber, 1939, Lloydia 2:146. M. hamulutu: Creighton 1950:99. M. hamulata hamulata: Smith 1979:1349. Record: GARFIELD: Rubys Inn (US). Creighton (1950) and Smith (1979:1349) list two forms of this species from midwest and western United States, including Utah, Colorado, and Arizona, nesting in upland plateaus between 7000 and 8000 ft. The Utah form is likely hamulata. Gregg (1963:300) lists this species between 8000 and 8700 ft in conifers, oak, and manzanita in Colorado. Myrmica incompleta Provancher M. incompleta Provancher, 1881, Nat. Canad. 12:359; Smith 1979:1349. M. rubra brevinodis var. sidcinodoides: Rees and Gnmd- mann 1940:5. M. rubra brevinodis: Rees and Grundmann 1940:5. M. brevinodis sulcinodoides: Cole 1942:368. M. brevinodis: Cole 1942:368; Creighton 1950:95; Ing- ham 1959:36; Knowlton 1970:211, 1975:6. M. incompleta incompleta: Smith 1979:1349. Records (Map 21): BOX ELDER: Kelton (K75) and 6 mi N (K70), Tremonton (KU). CACHE: Rlacksmith Fk Cyn, Elk Valley, W Hodges Cyn (KU), Logan (C42), Lo- gan Cyn (KU), Riverheights (RG), Tony Grove Cyn (KU). CARBON: Scofield (BAD). RICH: Garden City (KU), Laketown (BAD), PickleviUe (KU). SALT LAKE: Big Cottonwood Cyn, Salt Lake City (RG). SUMMIT: Woodland (C42). UINTAH: A.shley Crk nr Vernal (U). UTAH: Provo (BY). WASATCH: Deer Crk Res, Heber (U). WASHINGTON: Pine Valley (City) (BAD). WAYNE: Bicknell (U). WEBER: Beaver Crk head of, Monte Cristo 8 mi S, Roy 2 mi S (KU). COUNTY UN- KNOWN: Beaver Head (KU). Creighton (1950, as M. brevinodis) and Smith (1979:1349) list two races of this spe- cies, only one occurring in eastern to western United States, including Utah and Colorado, nesting under various objects through a wide elevational range. The Utah subspecies is likely sulcinodoides. Hunt and Snelling (1975:21) list this species from Arizona. Gregg (1963) lists it between 4600 and 10,500 ft under logs and rocks in juniper, ma- hogany, coniferous forest, grass, and other areas in Colorado. La Rivers (1968:2) lists it from Nevada, where Wheeler and Wheeler (1978:389) found it between 6400 and 9700 ft. Cole (1942:368) indicates its habitat in Utah as under stones, and Ingham (1959) found it under stones in southern Utah. In 27 recorded Utah localities it was found 11 times in montane areas. In 19 recorded elevations it was about equally distributed between 4225 and 7675 ft. Myrmica lobicornis Emery M. rubra scabrinodis var. fracticornis Emery, 1895, Zool. Jahrb. Syst. 8:313. M. scabrinodis lobocornis var. fracticornis: Rees and Grundmann 1940:5. M. lobicornis fracticornis: Cole 1942:368; Beck at al. 1967:71; Smith 1979:1349. M. lobicornis lobifrons: Creighton 1950:100; Ingham 1959:37; Knowlton 1975:6; Smith 1979:1350. M. lobifrons: Knowlton 1970:211. Records (Map 21): BOX ELDER: Snowville (K70). CACHE: Franklin Basin, Logan Cyn summit, Tony Grove Cvn (KU). DAGGETT: Radosovich Ranch (BAD). DUCHESNE: Fruitland (U), Neola (C42). GARFIELD: Boulder Mt (U). IRON: Cedar Breaks Nat Mon (159). KANE: AdairviUe (BAD), Long Valley Jet 11 mi W (1.59). RICH: Laketown (BAD). SALT LAKE: Alta, Big Cottonwood Cyn S fk (C42), Brighton, Butterfield Cyn, Red Butte Cyn (U). SANPETE: Bluebell Flats, Ephraim Cyn (KU), Wales (RG). SUMMIT: Camas (? = Kamas) (C42) and 21 mi E (A), Henrys Fk Basin (RG), Soapstone Cyn (RG). TOOELE: S Willow Cyn (U). UINTAH: Whiterocks Cyn (KU). UTAH: American Fk Cyn (U), Colton (BAD)^ WASATCH: Currant Crk (BAD), Deer Crk Res. (U). WASHINGTON: Pine Valley (City) (BAD). Creighton (1950) and Smith (1979) list two subspecies from eastern to western United States, including Utah, Colorado, and Ari- zona, nesting under stones or wood near 492 Great Basin Naturalist Vol. 42, No. 4 streams, where they are associated with Lep- tothorax provancheri. Records of both are re- ported from Utah. They may be separated by the antennal lamina, which encircles the bend of the scape in the form of a saucerlike flange on lobifrons but forms an angular toothlike projection on the inner side of the bend on fracticornis. Gregg (1963:303) lists this species between 6240 and 12,500 ft un- der rocks and logs in conifers, oak, manza- nita, birch, pinyon-juniper, sagebrush, and mahogany habitats in Colorado. La Rivers (1968:2) lists it from Nevada, where Wheeler and Wheeler (1978:389) found it between 6400 and 10,800 ft. They found it frequently in wood, and also commonly under rocks in North Dakota (1963). Allred and Cole (1971:239) found it in Idaho in association with a variety of shrub types. Cole (1942:368) indicates that it nests under stones and in logs in Utah. Ingham (1959) found it under stones or wood in coniferous forests in southern Utah. Knowlton (1975:6) found it associated with rabbitbrush in northern Utah. Twenty ants in one collection were taken from under a rock in conifers. In 31 reported Utah habitats it was taken 21 times from montane areas. In 19 recorded elevations be- tween 4544 and 10,500 ft it was about equal- ly distributed, taken once under 5000 and three times over 9000. Beck et al. (1967:71) found it feeding on dead rodents in six in- stances in Utah. Myrmica monticola Wheeler M. scabrinodis schenki var. monticola Wheeler, 1917, Proc. Amer. Acad. Arts Sci. Boston 52:505. M. scabrinodis: Hayward 1945:120. M. monticola: Knowlton 1970:211, 1975:7. Records (Map 21): BOX ELDER: Cedar Crk (City) (K70), Cedar Hill (K75), Curlew Valley (KU), Hardup (K75), Kelton, Kelton Pass, Snowville (K70), Wildcat Hills (K75). GRAND: Moab 10 mi SE (KU). SAN JUAN: Pack Crk (US). SANPETE: Majors Flat (KU). UTAH: Halls Fk rd 5.2 mi N Hobble Crk rd (A). COUNTY UN- KNOWN: locality given as "Mt Timpanogos or Uinta Mts" (H). Smith (1979:1350) lists this species from eastern to western United States, including Colorado, nesting under objects in wood- lands. Gregg (1963:309) lists it between 6000 and 8600 ft under rocks and logs in conifers, oak, pinyon-juniper, birch, and grass habitats in Colorado. Wheeler and Wheeler (1963) found it frequently in and under wood in 1 North Dakota. Knowlton (1975:7) found it as- sociated with sagebrush, grass, and junipers in northern Utah. One ant was collected from under a rock in a meadow association of grass, herbs, and sagebrush. Other ants under the same rock were M. emeryana, Formica fusca, and F. gnava. In 13 recorded Utah habitats it was taken only twice in montane forest. Two re- corded elevations were 4225 and 4544 ft. Neivamyrmx califomicus (Mayr) Eciton californicum: Mayr, 1870, Verh. Zool.-Bot. Ges. 20:969. Eciton sp.: Cole 1942:360; Ingham 1963:40. A^ califomicus: Smith 1979:1330. Records: "MILFORD CO" (C42) (? = Milford in Bea- ver Co). IRON: Modena (163). Smith (1979:1330) lists this species from western United States, including Utah and Nevada. Ingham (1963) found it in sagebrush in southern Utah. One recorded elevation in Utah was 5468 ft. Paratrechina parvula (Mayr) Prenolepis parvula Mayr, 1870, Zool.-Bot. Ges. Wien, Verh. 20:947. Paratrechina sp.: Beck et al. 1967:72. Paratrechina parvula: Smith 1979:1444. Records: DUCHESNE: Tabiona 11.6 mi E (A). SAN JUAN: Four Corners, Mexican Water (BAD). This is tentatively listed as parvula. Smith (1979:1444) lists it from eastern to western United States, including Utah and Arizona, nesting under stones and logs or in small cra- ters in open grassy areas. Sixteen ants in one collection in Utah were taken from under a rock in sagebrush. Beck et al. (1967:72) found it feeding on dead ro- dents in two instances in Utah. Pheidole bicarinata Forel P. bicarinata race vinelandica Forel, 1886, Ann. Soc. Ent. Belg. .30:45; Ingham 1963:78. P. bicarinata buccalis: Creighton 1950:171; Grundmann 1958:163. P. bicarinata: Ingham 1959:50; Beck et al. 1967:72. P. hngula: Ingham 1963:77. P. bicarinata paiute: Allred and Cole 1979:98. P. bicarinata vinelandica: Smith 1979:1.367. Records (Map 22): BEAVER: Frisco (BAD). BOX EL- DER: Lucin (BAD). CARBON: Wellington 10 mi NE (A). DUCHESNE: Roosevelt (BAD). GRAND: Dewey December 1982 Allred: Ants of Utah 493 Bridge, Moab (G58). KANE: Adairville, Cottonwood Crk (BAD), Glen Cvn City (AC), Navajo Wells (BAD). MILLARD: Desert Range' Exp Sta (BAD). SAN JUAN: Abajo Mts (U), Four Corners (BAD), La Sal Jet 1.6 and 23.1 mi S (A), Montezuma Crk (BAD). UINTAH: Jensen (BAD). WASHINGTON: Diamond Valley, Grafton (BAD), Pintura (159), Rockville, nr Short Crk (Arizona) (BAD), Snow Cvn (163), Springdale (159), Tinipoweap Cvn (163), Toquerville (BAD), Zion Nat Park (159). Creighton (1950) and Smith (1979) list four races of this species primarily as eastern and central United States, occurring in Colorado, Utah, Arizona, Nevada, and Wyoming, nest- ing in logs, exposed soil, or under objects in grassy areas. The subspecies may be sepa- rated by the following key. 1. Basal face of epinotum in the major largely covered with transverse striae bicarinata — Basal face of epinotum in the major largely punctate 2 2(1). Epinotum of minor with angular teeth, broad at the base and not resembling spines longula — Epinotum with thick, short spines 3 3(2). Color of major reddish to blackish brown, minor dull yellow to blackish brown paiute — Color of major clear yellow to yellowish brown, minor clear yellow vinelandica Gregg (1963:408) lists it in Colorado be- Creighton (1950) lists five and Smith tween 3500 and 6970 ft under rocks and logs (1979:1368) three races of this species from in a variety of habitats. Cole (1966:15) found western United States, including Idaho, Ari- it in southern Nevada under stones in various zona, and Nevada. The two races recorded types of desert brush. Wheeler and Wheeler for Utah may be separated by the occipital (1963) found it about equally under rocks and ^^gae of the major, which are coarse and wood in North Dakota. Ingham (1959, 1963) ^^vy on californica and fine and essentially found it in southern Utah under stones or in straight on oregonica. Cole (1942:362) in- roots of oak and rabbitbrush, or in craterlike dicates the habitat of this species in Utah as mounds in open areas in a variety of vegeta- ^^er stones in dry grassy areas. Grundmann tive types. Allred and Cole (1979:99) found it (1958:163) indicates that in Utah it nests un- in southern Utah in a wide variety of desert ^er stones or in soil around roots, generally in shrub types, abundantly in sagebrush. grass habitats. Ingham (1959, 1963: found it Twenty ants in one collection were taken i^ dry sandy soil in sagebrush in southern from a small mound, 50 in one collection Utah. Only 3 of 11 localities in Utah were in from under a rock, and one singly in an open montane forest. Twelve recorded elevations area. Three collections were taken in sage- ranged from 4042 to 7125 ft, 11 under 6000. brush: one in association with grass, one pin- yon, and one juniper and pinyon. Only one of Pheidole ceres Wheeler 27 localities in Utah was in a montane forest. r\( no J J 1 •-• u .- ncnn J P- ceres Wheeler, 1904, Bull. Amer. Mus. Nat. Hist. Of 23 recorded elevations between 2500 and on in n a in=:c icq n i * i ihctto r r>/^ nnr^^ 20: 10; Grundmann 1958: 163; Bcck et al. 1967:72. 7500 ft, 20 were between 3000 and 6000. Records: KANE: Kanab (G58). SUMMIT: Echo Beck et al. (1967:72) found it feeding on dead (BAD). rodents in 19 instances in Utah. Smith (1979:1368) lists this species as southwestern United States, including Colo- Pheidole californica Mayr rado and Arizona, nesting under stones in dry, sunny localities between 5000 and 9000 P. californica Mayr, 1870, Verh. Zool. Bot. Ges. Wien f^ Q^ggg (1963:413) lists it from Colorado 20:987; Rees and Grundmann 1940:4; Cole , , rA^-r j ocnr» r. j i • 1942:362; Grundmann 1958:163. between 5947 and 8500 ft under rocks m a P. californica oregonica: Cole 1942:362; Ingham 1959:52. variety of habitats, predominantly pinyon- Records (Map 22): GRAND: Moab (C42). KANE: Ka- juniper. In Utah it prefers plateaus between nab (C42). SALT LAKE: Big Cottonwood Cyn (U), Ft 5000 and 8000 ft and nests under stones in ^ATniiS'^Ah''' m/7S/\'^' ?^r'i^ Tnn^^i- desert conditions (Grundmann 1958:163). SAN JUAN: Abajo Mts (G58), La Sal (C42). TOOELE: ^ i i n • • n i Clover, Fisher Pass (C42). UTAH: Ironton (C42). Two recorded collections m Utah were at 494 Great Basin Naturalist Vol. 42, No. 4 4973 and 5467 ft. Beck et al. (1967:72) found it feeding on dead rodents in one instance in Utah. Pheidole dentata Mayr P. inorrisi var. dentata Mayr, 1886, Verb. Zool.-Bot. Ges. Wien 36:457. P. dentata: Beck et al. 1967:72. Records: GRAND: Dewey (U). WASHINGTON: To- querville (BAD). Smith (1979:1369) hsts this species from eastern to midwestern United States (no in- termountain state is hsted), where it nests in soil mounds or under various objects. Beck et al. (1967:72) found it feeding on dead rodents in one instance in Utah. Pheidole desertorwn Wheeler P. desertorwn Wheeler, 1906, Bull. Amer. Mus. Nat. Hist. 22:337; Cole 1942:362; Grundmann 1958:163; Ingham 19,59:52; Smith 1979:1369. Records (Map 22); GRAND: Moab (U). UINTAH: Red Cloud Loop rd 9.2 mi W U44 (A). WASATCH: Hanna 4.4 mi W (A). WASHINGTON: Castle Cliff (C42), Virgin City (159). Smith (1979:1369) lists this as a western species, including Utah, Arizona, and Ne- vada, nesting under stones and in small crater mounds. Cole (1966:15) found it in southern Nevada under large stones in various habitats of desert shrubs. Grundmann (1958:163) in- dicates that in Utah it lives under stones in desert habitats under 4500 ft. Ingham (1959, 1963) found it under rocks in blackbrush, yucca, mesquite, and four-wing saltbush in southern Utah. Seventy-three ants in two collections were found under rocks, once under the same rock with Formica argentea and F. pallidefulva. One collection was taken in a grassy mead- ow. In five recorded Utah localities two were in canyon-montane forest. Recorded eleva- tions range between 2500 and 4000 ft. Pheidole grundmanni M.R. Smith P. grundmanni Smith, 1953, J. New York Ent. Soc. 61:144; Smith 1979:1370. Record: UINTAH: Merkeley Park on Ashlet Crk (Sm53) (? = Ashley Crk). Smith (1979:1370) lists this species only from Utah. It was taken from under a stone in cottonwoods at 6000 ft (Smith 1953:144). Pheidole hyatti Emery P. hyatti Emery, 1895, Zool. Jahrb. Syst. 8:289; Ingham 1959:53; Beck et al. 1967:72. Records: IRON: New Harmony (1.59). KANE: Adair- ville (BAD). WASHINGTON: Grafton, RockviUe (BAD). Creighton (1950) and Smith (1979:1370) list two forms of this species from midwest to western United States, including Colorado and Nevada. The Utah form is likely hyatti. Hunt and Snelling (1975:21) list it from Ari- zona. Gregg (1963:419) lists it between 5500 and 5900 ft under rocks in grassy habitats in Colorado. Ingham (1959) found it under rocks in oak, pinyon-juniper, rabbitbrush, and sagebrush in southern Utah. None of four recorded Utah habitats were in montane areas. Four recorded elevations were between 3660 and 6000 ft. Beck et al. (1967:72) found it feeding on dead rodents in three instances in Utah. Pheidole pilifera Cole P. pilifera artemisia Cole, 1933, Ann. Ent. Soc. Amer. 26:616; Rees and Grundmann 1940:4; Cole 1942:,362; Creighton 1950:187; Ingham 1959:53; Smith 1979:1372. Records (Map 22): EMERY: Hideout Cyn nr Green River (U). GARFIELD: Boulder (U). SALT LAKE: Big Cottonwood Cyn (U). SEVIER: Salina Cvn nr Fremont Jet (U). UINTAH: Gusher (C42). UTAH:' Payson (C42), Provo (RG), Provo Cvn (U), Wanrhodes Cyn (KU). WASHINGTON: Springdale (C42). Creighton (1950) and Smith (1979:1372) list four races of this species from eastern to western United States, including Utah, Colo- rado, Arizona, and Nevada, nesting under stones or in craterlike excavations in open areas where they are associated with P. in- quilina. The two subspecies that likely occur in Utah may be separated by the surface of the occiput of the major, which has promi- nent rugae on artemisia, whereas on colora- densis the surface is finely and contiguously punctate. Gregg (1963:422) lists this species from Colorado between 5100 and 8500 ft un- der rocks in a variety of habitats, pre- dominantly in pinyon-juniper. Cole (1966:15) found it in southern Nevada under large stones in pinyon-juniper. He indicates its hab- itat in Utah as sagebrush (1942:362). Wheeler and Wheeler (1963) found it frequently under rocks in North Dakota. Ingham (1959) found it under rocks in sagebrush in southern Utah. In 10 recorded Utah habitats it was taken 3 December 1982 Allred: Ants of Utah 495 times in montane forest. In 11 recorded ele- vations it was about equally distributed be- tween 3900 and 7000 ft, twice over 7000. Pheidole sitarches Wheeler P. soritis Wheeler, 1908, Bull. Amer. Mus. Nat. Hist. 24:439. P. sitarches soritis: Creighton 1950:190; Cole 1956:116; Ingham 1959:53; Allred and Cole 1979:99; Smith 1979:1373. Records (Map 23): KANE: Glen Cyn City (AC). UIN- TAH: Dry Fk rd 13.4 mi N U121 (A). UTAH: Provo (C56). WASATCH: Hanna 9.2 mi W (A). WASHING- TON: nr Short Crk (Arizona) (159). COUNTY UN- KNOWN: Skiill Valley (? probably Tooele Co) (C56). Creighton (1950) lists three and Smith (1979:1373) four races of this species from eastern to western United States, including Utah, Colorado, and Arizona. The Utah sub- species is likely soritis, which may be sepa- rated from the close Colorado form by the head of the minor, which is striate posteriorly on soritis but pimctate on campestris. Gregg (1963:424) lists this species between 3500 and 4400 ft under rocks in cottonwood-willow and grassy habitats in Colorado. Ingham (1959, 1963) found it in low mounds in bur sage, little rabbitbrush, junipers, and sage- brush in southern Utah. Allred and Cole (1979:99) found it in southern Utah in a vari- ety of desert shrub types, most frequently in ephedra-grass. Sixty ants in two collections were found under rocks. One was in sagebrush and one was in sagebrush in association with snow- berry. In six recorded Utah habitats it was taken twice from montane forest. Three re- corded elevations were between 3250 and 5000 ft. Pheidole virago Wheeler P. virago Wheeler, 1915, Amer. Mus. Nat. Hist., Bull. 34:401; Ingham 1959:54. Record: WASHINGTON: between Hurricane and Harrisburg Jet (159). Smith (1979:1374) lists this species from Texas and Arizona. Ingham (1959, 1963) found it in southern Utah in small crater mounds in creosote bush, bur sage, little rab- bitbrush, cholla, marigold, and Russian thistle. In two recorded Utah collections it was taken at 2900 and 3000 ft. Pogonomyrmex barbatus (M.R. Smith) Mynnica hurbata Smith, 1858, Cat. Hym. Brit. Mus. 6:130. P. barbatus molefaciens: Olsen 1934:501; Rees and Grundmann 1940:4; Cole 1942:367. P. barbatus tiiarfensis: Rees and Grundmann 1940:4; Cole 1942:367. P. barbatus fuscatus: Rees and Grundmann 1940:4; Cole 1942:367. Records: EMERY: Goblin Valley 10 mi E (KU). WASHINGTON: St George (O). WAYNE: Fruita 5 mi SE (U). Smith (1979:1353) lists the distribution of this species as eastern United States westward to include Colorado, Arizona, and Nevada, nesting in low to high craterlike mounds. Grundmann (1958:162) designated it as a southern Utah and northern Arizona desert form, nesting in low craterlike mounds in sandy soil. Cole (1968:56) believes that the St. George (Washington County) record for Utah listed by Olsen (1934) may represent a sparse population of short duration. Cole's distribution map (p. 57) shows it considerably east and south of Utah. Pogonomyrmex brevispinosus Cole P. brevispinosus Cole, 1968, Univ. Tenn. Press, p. 89. Record: KANE: Wahweap (KU). Smith (1979:1354) lists this species from western United States, including Nevada, nesting in low crater mounds. Its occurrence in Utah is questionable. Pogonomyrmex californicus (Buckley) Myrmica californica Buckley, 1867, Proc. Ent. Soc. Phila. 6:336. P. californicus: Olsen 1934:502; Rees and Grundmann 1940:5; Cole 1942:367, 1968:120; Creighton 1950:123; Grundmann 1958:162; Ingham 1959:41; Smith 1979:1356. Records (Map 23): KANE: Glen Cyn Res, Kanab (G58). SAN JUAN: Hole-in-the-Rock Cyn (U). WASH- INGTON: Beaver Dam Wash (159), Hurricane (RG), be- tween Hurricane and Harrisburg Jet (159), Leeds, Leeds Cyn (KU), Santa Clara (US), Santa Clara Crk (KU), Shiv- w'its Indian Res (159), Snow Cvn (KU), St George (O), Veyo (159), Virgin, Washington (KU). Smith (1979:1356) indicates the range of this species as primarily southwestern United States, including Arizona and Nevada, in cir- cular or semicircular craters of loose sand. Cole (1966:4) states that in southern Nevada* its nests are craterlike, occurring principall) in hopsage-matrimony vine and Russian 496 Great Basin Naturalist Vol. 42, No. 4 thistle habitats. He (1968:121) shows it in ex- treme southern parts of Utah essentially west of the Colorado River and in much of Ari- zona and Nevada. Gnindmann (1958:162) designates its distribution in Utah as below 4000 ft, basically a Lower Sonoran form. Ing- ham (1959) indicates its habitat in southern Utah as conical mounds in creosote bush, sagebrush, and juniper. None of its Utah col- lections were in canyon-montane areas. In 16 recorded elevations it was between 2500 and 4973 ft, 9 under 3000. Pogonomyrmex imberbicuhis Wheeler P. imberbicidm Wheeler, 1902, Amer. Nat. 36:87; Ing- ham 1959:46; AUred and Cole 1979:99. Records: KANE: Glen Cvn City (AC). WASHING- TON: Hurricane (159). Smith (1979:1357) lists this species from midwest to western United States, including Colorado, Arizona, and Nevada, nesting un- der stones or in small craters in open areas. Gregg (1963:336) lists it at 4800 ft in saltbush habitats in Colorado. Cole (1968:168) shows it in isolated collections in southeastern Ne- vada and Arizona only in the extreme south- eastern comer. He states that in southern Ne- vada it nests under stones in hopsage and matrimony vine (1966:6). Ingham (1959, 1963) designates its habitat in southern Utah as soil without mounds in creosote bush, blackbnish, )TJCca, choUa cactus, and bur sage. AUred and Cole (1979:99) found it in a saltbush-sagebrush association in southern Utah. Three Utah collections were between 3250 and 3500 ft. Pogonomyrmex maricopa Wheeler P. californicus maricopa Wheeler, 1914, Psyche 21:155. P. maricopa: Ingham 1963:47; Cole 1968:138; Smith 1979:1356. Records (Map 23): JUAB: Chicken Crk Res (KU). MILLARD: Sutherland (US). WASHINGTON: Hurri- cane (BY), Leeds, Leeds Cyn (US), Santa Clara (KU), St George (BY), Washington (KU), Smith (1979:1356) lists this species from western United States, including Utah, Colo- rado, Arizona, and Nevada, nesting in crater- like mounds. Gregg (1963:327) lists it at 4600 ft in weedy areas in Colorado. Cole (1968:139) shows it across extreme southern Utah extending northward along the Colo- rado River drainage and southward into much of Arizona, as well as southwestern Ne- vada. Ingham (1959, 1963) found it in sand sagebrush and little rabbitbrush in southern Utah. Eight recorded Utah habitats were in desert areas between 2500 and 5000 ft, most- ly at lower elevations. Its occurrence in Juab and Millard counties is questionable; such records likely are occidentalis. Pogonomyrmex occidentalis (Cresson) Mi/rmica occidentalis Cresson, 1865, Proc. Ent. Soc. Phila. 4:426. P. occidentalis: Olsen 1934:507, 509; Rees and Grund- mann 1940:5; Cole 1942:365; 1968:94; Beck et al. 1967:72; Knowlton 1970:211, 1975:7; Allred and Cole 1979:99; Smith 1979:1355. P. occidentalis iitahensis: Olsen 1934:509; Cole 1942:.365. P. occidentalis comanche: Creighton 1950:128; Grund- mann 1958:162; Ingham 1959:44. P. occidentalis occidentalis: Grundmann 1958:162. Records (Map 24): BEAVER: Beaver (RG), Lund 35 and 49 mi N, Milford 1.3 mi N, and 11.5 and 26.7 mi W (A), Wildcat Cvn (C42). BOX ELDER: Blue Crk (City), Bovine (C42), Brigham (O) and 0.9 mi E (A), Collinston (C42), Corinne (US) and 6.1 and 16.1 mi W, Deweyville 0.3 mi N (A), Garland, Hansel (US), Hansel Mts (K70), Hardup, Hov^/ell (C42), Kelton 17.2 and 28.5 mi SW (A), Lampo (US), Locomotive Spngs, Lucin (C42), Promon- tory 1.3 and 10.3 mi W (A), Promontory Pt (C42), Snow- ville (O) and 7.7, 19.1 and 30.7 mi S (A), Thicket (KU), Wendover 28 mi N (A), Willard (C42) and 1.5 mi S (A). CACHE: Blacksmith Fk Cyn (KU), Cache Jet (C42), Cornish (KU), Cove (C42), Leeds Cyn (KU), Logan (C42), Logan Cyn (KU), Mendon, Providence (C42), Smithfield 0.5 m'i N (A), Mt Sterling (C42). CARBON: .\rgyle Cyn, Kenilworth (U), Myton rd 5 and 22.7 mi E US6, Price 2.2 mi S (A) and 20 mi S (WU), Wellington 5 mi S, and 5 and 10 mi NE (A). DAGGETT: Radosovich Ranch (BAD). DAVIS: Farmington (C42), Kaysville (O), Layton 1.5 mi S (A). DUCHESNE: Blue Bench, Currant Crk (C42), Duchesne 5.3 mi E, 2 mi N (A) and 11 mi W (WU), Hanna 1.5 mi W, Myton rd .33.5 mi E US6, Neola 6.2 mi S, Tabiona 11.6 mi E, Wellington rd 7.2, 17.3 and 27.4 mi S US40 (A). EMERY: Buckhorn Res (BY), Emery 4.5 mi N and 4.9 mi S, Ferron 1.4 mi N (A), Goblin Val- ley and 10 mi E (KU), Green River (C42), Hanksville 16 mi N (KU), Hideout Cyn nr Green River (U), Hunting- ton (US) and 1.7 mi S, Orangeville 3 mi W, Wellington 15, 26, .36 and 46 mi S (A). GARFIELD: Antimony 5.8 and 27.6 mi S (A), Boulder Mts, Carcass Crk (on Boulder Mt) (U), Bryce Cyn Nat Park (US), Escalante 20 mi S and 20 mi E (U), Henry Mts (G58), Hanksville 26.6 mi S (A), Osiris (U), Rubys Inn (US), Tropic 3.3 mi E, Jet U12 and U63 0.5 mi E (A). GRAND: Arches Nat Park, Cisco (BY) and 1.5 mi E, Crescent Jet 5 and 18.8 mi S, and 6, 37 and 47 mi E (A), Dewey (U), Green River 5 and 16 mi E (A), Moab (G58) and l'.5 mi S (A), Thompson (C42). IRON: Beryl 1 mi NE (A), Cedar City (RAU), 3.7 mi E, and 6.3, 16.8 and 27 mi W (A), Columbia Iron Mine 7 mi W (159), Enoch (RAU), Iron Mt (US), Little Pinto December 1982 Allred: Ants of Utah 497 (KU) and 3 mi NE (US), Lund 8.6 and 19 mi N, Modena 5.3 and 8.7 mi W, and 10.4 and 25 mi NE, Newcastle 7.2 and 17.2 mi W (A). JUAB: Callao 7.8 mi E, Eureka 0.5 mi E and 3.7 mi S (A), Fish Spngs (BY), 0.8 mi N, and 1, 10.6, 20.8 and 30.8 mi E, Gandy 10.1 mi N (A), Indian Farm Cyn (in Deep Crk Mts) (U), Jericho (BY), Levan 6.9 mi N and 10 mi S, Lvnndyl 11.5 and 21.5 mi N, Nephi 3 mi N, and 5.5, 12.1 and 23 mi W (A), Pony Ex- press Sta (US), Tintic (C42), Topaz Mt and 5 mi NW (US), Trout Crk (City) 2.8 mi N (A). KANE: Adairville (BAD), Cannonville 8.7, 31.3 and 41 mi S (A), Escalante River (BY), Glen Cyn City (AC), Johnson Cyn (BY), Ka- nab (O), 4-7 mi N (159), 4 mi E (KU), and 5, 20.4, 31 and 41 mi E (A), Long Valley Jet 2 mi W (159) and 0.5 mi S, Mt Carmel Jet, Orderviile 2 mi N (A), Paria (BY), Wah- weep Cmpgnd (US). MILLARD: Black Rock (KU), Del- ta (BY) and'^0.5 mi W (A), Deseret (US), 3 mi S (KU) and 30 mi S (US), Fillmore (C42), Hinckley 4.8, 14.7, 24.9, .35, 45, 58.1 and 69.6 mi W (A), Holden, Kanosh (US), Lvnndyl 2 mi S (A), Meadow (C42), Milford 16.3, 31.5, 46.3 and 61.3 mi N (A). PIUTE: Antimony 4.8 mi N (A), Grass Valley (KU), Kingston (A), Marysvale (BY) and 4.1 mi S (A). RICH: Garden City 1.6 mi N, Randolph 2.3 and 5.3 mi N, Woodruff and 4 mi W (A). SALT LAKE: Big Cottonwood Cyn, Ft Douglas, Granite, Point-of-Mt (C42), Riverton 2.6 mi S (A), Salt Lake City (C42), Sandy (O). SAN JUAN: Abajo Mts (G58), Blanding 6.7 mi N (A) and 8 mi N (U), Blue Crk, Bluff (C42) and 0.6 and 11.6 mi S (A), Jet Colorado and San Juan rivers (LI), La Sal, La Sal Jet (RG) and 1.6, 11.7 and 23.1 mi S, Mexican Hat 8.2 mi S, Moab 12 mi S, Monticello 0.5 mi N, Jet U95 and U261 7.7 mi W, Jet U261 and US163 9.4, 19.6 and 29.9 mi N (A), Squaw Flat Cmpgnd (in Canyonlands Nat Park) (WU). SANPETE: Ephraim and 3 mi N (A), Eph- raim Cyn (U), Fairview (BY) and 16.5 mi N, Freedom 0.5 mi N, Gunnison (A), Indianola (BY), Levan 20 mi S (A), Majors Flats (KU), Mt Pleasant (C42) and 0.5 mi N, Nebo Loop rd 6.3 mi E Jet Ull (A), Wales (RG). SE- VIER: Axtel 4 mi S, Big Rock Candy Mt (A), Elsinore (C42), Fremont 18.9 and 28.6 mi N, Richfield 2.3 and 7.5 mi S (A). SUMMIT: Coalville (U), Francis 1 mi E, Wyoming brdr 1.5 and 10.4 mi W on 180 (A). TOOELE: Clover 1 mi W (A), Delle (O) and 2, 12 and 22 mi W (A), Dugway (BY) and 5 and 10 mi E, Faust 1.5 mi W and 3.6 mi E (A), Grantsville (O) and 2 and 12.1 mi W (A), losepa (C42), Johnson Pass (US), Little Valley (C42), Lookout Pass (BY), Low, Orrs Ranch (C42), Rowley Jet 0.5, 10.8, 21 and 31 mi S, Simpson Spngs 4 and 14 mi E (A), Stansbury Island (C42), Tooele (O), 1 mi S and 3.3 mi W, Jet U36 and U73, Vernon, Wendover 10.3, 13.3, 14.5 and 23.1 mi N (A), S Willow Cyn (U). UINTAH: Bonanza 3 mi S (KLI), 8 and 21 mi S, and 5 and 15 mi N, Dry Fk rd 9.4 mi N U121 (A), Ft Duchesne (C42), Gush- er 4 mi E (KU), Jensen 5.5 mi W and 7.5 mi E, Lapoint 1 and 10.7 mi E (A), Ouray (C42), Split Mt, Vernal (BY), 6 mi E (KU) and 5 and lo'mi N (A). UTAH: Cedar Fort (U) and 1.9 mi N (A), Elberta (C42) and 1 mi W, Fair- field 3.3 mi W (A), Lehi (O) and 4.4 mi W (A), Orem, Provo, Provo Cyn (BY), Santaquin (C42), 1.5 mi N, 1.5 mi W and 0.5 mi S, Springville 2 mi S, Thistle 2.7, 3.6, 7.7, 9 and 14.6 mi E (A), W Utah Lake (BY). WASATCH: Francis 5.4 mi W, Hanna 4.4 mi W (A), Heber (U) and 2.3 mi W (A), Soapstone Cyn (U). WASH- INGTON: Grafton (159), Hurricane (RAU)^ Leeds (RG), Mt Meadow, New Harmony (BY), Pine Valley (City) (159), Pinto, 3 mi S (US) and 5 mi S (KU), Rockville, San- ta Clara (159), Springdale (C42), St George (RG), Virgin, Zion Nat Park (O). WAYNE: Capitol Reef Nat Park, Fruita 5 mi SE (U), Hanksville 3, 17.4 and 27.2 mi W, and 7 and 16.7 mi S, Torrey 3.7 mi E and 5.3 mi W (A). WEBER: Little Mt (C42), Ogden (O), Ogden Cyn (BY), Riverdale (C42), Jet U39 and U166 3.3 mi E, Woodruff 34.8 mi W (A). COUNTY UNKNOWN: Brush Crk (RG) (prob Uintah Co), Westville (C42) (prob Wellsville in Cache Co). Smith (1979:1355) indicates this species as a western one, inckiding Utah, Colorado, Ari- zona, Nevada, Idaho, and Wyoming. Gregg (1963:331) hsts it from Colorado between 3500 and 9000 ft in a variety of habitats, pre- dominantly in pinyon-juniper, sagebrush, and grass. Ingham (1959, 1963) indicates its habi- tat in southern Utah as a variety of shrub types. Allred and Cole (1979:99) found it in southern Utah in a wide variety of desert shrub types, most frequently in grass-hopsage and ephedra-grass-blackbrush-hopsage associ- ations. Rees and Gnmdmann (1940:5), Cole (1942:365), and Grundmann (1958:162) list it from a variety of situations in Utah. Thirty-two hundred ants in 211 collections were taken from 87 different plant types or associations. Eighty-one collections were taken in sagebrush or sage associations. Fifty- two collections were in grass or grass associ- ations other than with sagebrush. Twelve col- lections were from pure stands of grease- wood, and 10 collections from greasewood with other plants except sagebrush or grass. Seventeen collections were from halogeton associated with other plants except sagebrush or grass. Thirteen collections were from shadscale in association with other plants ex- cept sagebrush or grass. Only 8 collections were taken from rabbitbrush associations other than with sagebrush or grass, and only 5 collections from blackbrush. Only 2 collec- tions were taken in juniper and/or pinyon where other plants were not present, but 19 collections where sagebrush was associated. This species commonly inhabits the shoul- ders and barrow pits along roads. Whenever sedges, meadow grass, or salt grass were pres- ent alongside the roads, I never found colo- nies of this species. Apparently it does not like the saline environment in this type of sit- uation. Colonies were seldom found in heavy clay soils, but occasionally in lighter clays. 498 Great Basin Naturalist Vol. 42, No. 4 Unstable sandy soil does not support this spe- cies, apparently because of the shifting po- tential of the sand from frequent winds. From just west of Hinckley in Millard County for a distance of about 14 miles westward, the soil is alkaline clay with greasewood, saltbush, and shadscale. In this area harvester mounds were found only along the shoulder of the highway where a gravel substrate was pres- ent. None were found on the alkaline flats until the greasewood and saltbush were re- placed by shadscale and desert pavement. Be- tween Sagecreek Junction and Laketown in Rich County, a distance of 10 miles where the soil is a reddish clay loam, not a single harvester mound was seen. Typical mounds were not seen between Blanding and Bluff in San Juan County, where the elevation dropped from 6100 to 4300 ft in 70 miles, and only those of P. rugosus were present. South of Bluff past Mexican Hat both mound forms and species were present, but occiden- talis was not as common as in other areas. About 11 miles north of Mexican Hat follow- ing highway U261, Cedar Mesa lies at 6500 ft. Mounds of rugosus were found up to its base but were replaced on top by occiden- talis. Ants of rugosus occur infrequently fur- ther westward along U95, where the eleva- tion drops back to under 4600 ft. In the 375 localities listed, only 31 were in montane forest. The 119 recorded elevations varied from 2750 to 9000 ft, 86 of them be- tween 4000 and 6000. Only 21 were over 6000 ft, 2 over 8000. There were 213 typical mounds observed. In 5 cases two main mounds were joined and apparently inhabited by the same colony. In only one instance did I find a colony oc- cupying three joined mounds. In a grease- wood habitat 17.2 mi southwest of Kelton in Box Elder County a colony was found with three joined, small linear mounds that had a total of nine entrances. A typical mound in an alkali-greasewood flat 7.8 mi east of Cal- lao in Juab County was covered with black chips of desert pavement. In an area 4.9 mi south of Emery along UIO at 6000 ft in Emery County in greasewood, grass, and Rus- sian thistle several mounds were completely covered with a layer of black gravel, which undoubtedly was carried from the shoulder of the road where it had been used for sur- facing. Some such mounds were as far as 60 ft from the road. Four mounds on an east-fac- ing slope at the north edge of Woodruff at 6400 ft in Rich County in sagebrush were covered with large gravel, the particles much larger than seen on other harvester mounds elsewhere in the state. These also were elon- gate and ridged, not the symmetrical cones like the typical harvester mounds. A low mound was found two feet from the asphalt 14.7 miles west of Hinckley in Millard Coun- ty, with the opening at the base on the north- east side. The area around the entrance was coated with a hardened amber material re- sembling baltic amber or shellac. Apparently this was deposited from the feet of the work- ers over a long period of time, perhaps picked up as a distillate from the hot surface of the adjacent asphalt. No other situation of this type was observed with any other mound over the state. A mound in another area had two rodent burrows in it, with a gopher snake curled in one of them. Food storage of seeds was frequently found in the upper part of the mounds, usually within a few inches of the top. In northwestern Utah around Kelton in Box Elder County, where the range of three species of harvesters overlaps, occidentalis are much larger than most salinus, but about the same size as owyheei. To see the angled basal tooth on the man- dible, one sometimes must study a series for best observation of open mandibles. On some specimens all teeth of both mandibles are well developed and the basal tooth on both sides is well developed and distinctly angled. On other specimens the basal tooth on one side may be vestigial, in which case the other teeth on the same mandible may be much re- duced, almost lacking in some cases, or mod- erately well developed in others. The basal tooth on the other mandible, however, is well developed and distinctly angled. Frequently the angle is not distinct as such, but the basal tooth forms a gentle curve instead. In series 388, taken 3.3 miles west of Tooele alongside highway U112 in Tooele County, one ant has eight teeth on the right mandible and seven on the left. The basal tooth is offset on both mandibles. In series 122, taken 37 miles east of Crescent Junction alongside highway 170 December 1982 Allred: Ants of Utah 499 in Grand County, most specimens have a bas- al tooth that is only slightly angled with the posterior border of the mandible, forming al- most an equal curvature. In series 305, taken five miles south of Crescent Junction along- side highway US 163 in Grand County, most specimens have a basal tooth that is not as offset as most other typical occidentalis scat- tered throughout Utah. On some of these specimens one side of the mandible is almost straight, whereas on the other, or on both sides on some specimens, the basal tooth forms a slight gentle curve. Other characters, such as the rugae and puncta on the head and the shape of the base of the antenna, are typ- ical characters. In series 113, taken 26 miles south of Wellington alongside highway US6 in Emery County, the basal tooth on both mandibles is well developed but the posterior margin of the mandible is almost straight without an angle. A colony (series 115), taken 46 miles south of Wellington in Emery Coun- ty in halogeton in mid-July, consisted of many workers with a few immatures but no winged forms. In a series of 30 workers exam- ined, the basal tooth on many is almost in a straight line with the basal margin of the mandible, and on a few it is in straight align- ment. Other characteristics are typical of oc- cidentalis. In series 199, taken one mile east of Francis in Summit County the basal tooth is often reduced on one mandible. On some specimens where it is reduced, the border of the reduced tooth is not at a distinct angle, but almost straight. In series 354, taken 17.4 miles west of Hanksville on highway U24 in Wayne County, many workers have essen- tially a straight mandibular border; others have an angled basal tooth at least on one side, but not as drastically as typical occiden- talis of other areas of Utah. On some workers all of the teeth on both mandibles except the two basal ones are worn off so that the inside edge of the man- dible is essentially straight except for the two basal teeth, which are typically offset. In series 356, taken 3.7 miles east of Torrey in Wayne County, one worker has both man- dibles with no evidence of any teeth. The edge is heavily chitinized, but it appears that all the teeth have been worn off smooth so that the normal toothed edge is perfectly straight, with no indentation showing where teeth should be, even to the offset basal tooth, which is also missing on both sides. Some intergradation of mandibles occurs in northwestern Utah, mainly between salinus and owyheei. Most occidentalis in that area are typical, but some have atypical man- dibles close to salinus and owyheei. In series 549, taken 17.2 miles southwest of Kelton in Box Elder County, some ants have typical oc- cidentalis mandibles, antennae, and petiole pattern; some have mandibles and antennae of occidentalis, with a petiole pattern of sa- linus; some have mandibles of salinus or owyheei and antennae of occidentalis, but with a petiole typical of salinus. In 111 mounds where other than workers were found, immatures but no winged forms were found in 20 (18 percent), winged forms only in 47 (42 percent), and both immatures and winged in 44 (40 percent). Immatures and winged forms were found in mounds above normal ground level only between 28 June and 3 August. When immatures were found in the summer they were between two to four inches below the outer surface of the mound in different parts of the mound as ori- ented to the direction of the sun. This may be optimum for brooding temperatures of the immatures, and they likely are moved period- ically by workers as the outside temperature and rays of the sun change. On 31 July in sagebrush one mile east of Brigham in Box Elder County at 0650, a mound was observed that was still in shade. No ants were outside the mound, but many winged forms were congregated just inside the entrance, probably preparatory for the nuptial flight. Almost as many winged ants as workers were found inside the mound. On the same day in a grass and sunflower habitat 6.1 miles west of Corinne in Box Elder Coun- ty at 0740, a small mound was observed that was still in shadow. Many winged forms were assembled just inside the entrance, but no ants were outside. The same situation was seen in two other adjacent mounds. In a mound located three miles west of Orange- ville at 6000 ft in Emery County in rabbit- brush, greasewood, and herbs near an irri- gated farm, winged forms were much more numerous than workers on 31 July and were found near the opening to the nest, whereas the fewer workers were deeper in the mound. 500 Great Basin Naturalist Vol. 42, No. 4 Apparently tlie winged forms were nearing their swarming time. Winged forms vary in depth in the mound, but frequently can be seen "peeking" from the openings. Callow (newly emerged) work- ers frequently were found mixed with the winged forms and were the principal workers to rescue immature stages when the nest was disturbed. Winged females are much more active in trying to hide or escape when dis- turbed than are winged males. The winged females frequently aid the callow workers in rescuing the immature forms, but I never once saw a winged male performing that task. One colony taken 41 miles east of Kanab in Kane County was in a low mound in rabbit- brush, matchbrush, and saltbush. Moderate numbers of workers were present, but no im- mature or winged forms. These ants behaved differently than typical occidentalis— they were not aggressive and did not swarm to- ward the intruder when disturbed, but simply scurried around almost at random. Their ab- domens were darker (almost blackish) than the amber unicolor of typical occidentalis. Forty-five miles west of Hinckley, alongside highway US6-50 in Millard County, a colony was found in a typical occidentalis mound with the opening at the base on the southeast side. These ants behaved similarly to salinus. Tliey were not aggressive but tried to hide when disturbed. A mound covered with thick detritus and occupied by occidentalis was found 7.7 miles west of junction U261 on U95 in San Juan County in sagebrush, pinyon, and jimiper. Ants of Conomijnna insana were abundant on the mound, crawling on and be- tween the sticks. No nest openings of these small ants could be seen, nor were any of their nests found in the vicinity. No inter- action between the two species was observed as they intermingled on the mound. A large, flattened moimd was found in rabbitbrush, sagebrush, and juniper 19 miles north of Lund on the road to Pine Valley in Iron County in late July at 6700 ft. Workers were transporting sand particles from its opening. Many workers, immature, and winged forms were found in the mound. Nine small mounds occupied by Conomxjrma insana were on the harvester mound. No apparent competition or interaction was seen between the two spe- cies as they intermingled in their scurryings. Beck et al. (1967:72) found occidentalis feeding on dead rodents in only two instances in Utah. Pogonomynnex owyheei Cole P. occidentcilis owyheei Cole, 1938, Amer. Midi. Nat. 19:240. P. owyheei: Cole 1968:102; Knowlton 1970:211, 1975:7; Smith 1979:1355. Records (Map 24): BOX ELDER: Cedar Crk (City) (K75), Cedar Hill (US), Curlew Valley (K70), Hansel Mts, Hardup, Kelton (K75), 6 mi N (US), 8.3 mi E, 70.4 mi SW (A), and 12 mi NW (KU), Kelton Pass (K75), Kel- ton rd 5 mi E U30 (A), Locomotive Spngs (K75), Park Valley (City) 10 mi E (KU), Snowville (K75) and 1 mi W, Jet U30 and U42 and 3.8, 5.8, 6.1 and 6.7 mi N (A), Wildcat Hills (K75). EMERY: Goblin Vallev, Hanksville 16 mi N (KU). GRAND: Moab, Thompson (KU). JUAB: Fish Spngs (KU). MILLARD; Black Rock (KU). WASH- INGTON: Rockville, Santa Clara Crk, St George, Vir- gin, Zion Nat Park (KU). Smith (1979:1355) lists this species as west- ern United States, including Utah, Nevada, Idaho, and Wyoming, nesting in earthen mounds. Allred and Cole (1971:239) found it in Idaho abundantly in associations of rabbit- brush-sagebrush-grass, less commonly in other shrub associations. Knowlton (1975:7) found it associated with sagebrush and rab- bitbrush in northern Utah. There were 260 ants taken from typical conical mounds in nine collections. Imma- tures and winged forms were found in four mounds in late July. In two of the mounds only female alates were found. Seven collec- tions were taken in sagebrush: two in associ- ation with grass and one with grass and rab- bitbrush. Two collections were taken in greasewood. In a greasewood area alongside an old railroad bed 8.3 miles east of Kelton in Box Elder County, a colony was adjacent to several colonies of salinus. Elevations of owyheei in Utah are between 4310 and 5600 feet in desert habitats. On 31 July in Box Elder County two mounds were seen wherein many winged forms were concentrated just inside the en- trance. At one mound several winged females came out of the entrance but quickly re- entered when I approached. In series 558, taken 5.8 miles north of highway U30 along U42 in Box Elder County, the ants were not as aggressive as occidentalis nor as prone to December 1982 Allred: Ants of Utah 501 hide as solinus, but their behavior was closer to that of salinus. Fewer could be enticed to come out of the entrance than is the case with occidentalis when air is blown into the entrance. Workers of occidentalis literally swarm out in such a reaction. In series 560, taken 6.7 miles north of the junction of highways U30 and U42 alongside U42 in Box Elder County, many specimens have a broadly rounded thorax, not flattened. The superior lobe of the scape base is some- what variable. On several specimens on one side the lobe has a collar that is extended su- periorly, but with no up-curvature as on typi- cal occidentalis. On the other side of the same specimens the lobe is similar to salinus, angvilar and rounded, without the collar ex- tension. Both segments of the petiole lack the typical salinus transverse striae, although faint subparallel striae are on the posterior half of the petiole, whereas the postpetiole lacks such striae on a few specimens. Most specimens in other series have the superior lobe of both antennae with an extension of the collar. In series 559, taken 0.6 mile south- east of series 560 above, at least one speci- men has a superior lobe of the scape base that is almost typically salinus— rounded without the collar extension. On the other an- tenna the superior lobe is angled and rounded without the collar extension. In series 557, which was taken six miles southeast of the above collections, one specimen has typical oivijheei antennae and mandibles with an atypical salinus petiole. The records of this species from Juab, Mil- lard, Emery, Grand, and Washington coun- ties most likely are errors of identification of aberrant specimens of occidentalis, wherein the offset basal tooth of the mandible is not typical. Such variations were discussed under occidentalis previously. Pogonomijrmex rugosus Emery P. barbatus rugosus Emery, 1895, Zool. Jahrb. Syst. 8:309; Rees and Grundmann 1940:5; Cole 1942:366; Crcighton 1950:120; Grundmann 1958:162; Ingham 1959:39. P. rugosus: Cole 1968:70; Allred and Cole 1979:99; Smith 1979:1354. Records (Map 23): EMERY: Green River (RG). GRAND: Arches Nat Park (BY), Dewey (U), Green Riv- er S of (KU), Moab (U), Thompson (C42). KANE: Glen Cvn City (AC), Kanab (G58) and 4 and 40 mi E, Wah- weap (KU). SAN JUAN: Abajo Mts (U), Blanding 6.4 and 12.2 mi S (A), Bluff (KU) and 0.6 and 11.6 mi S l\). Four Corners (US), Goulding Mt (RG), Mexican Hat (U), Mon- ument Valley (KU), Navajo Bridge (on Highway 89) (G58), Jet U95 and U261 35 and 40.4 mi W, Jet 'U261 and US163 3.7 mi N (A). UTAH: Silver Lake Flat (A). WASHINGTON: Hurricane (RG), Harrisburg Jet 2 mi \V (159), Leeds Cyn, Rockville, Santa Clara, Snow Cyn (KU), Springdale (WU), St George (O), Virgin City, Zion Nat Park (159). Smith (1979:1354) lists this species from midwest to western United States, including Utah, Colorado, Arizona, and Nevada, nest- ing in the soil in a flattened gravel disc. Gregg (1963:323, 327) lists it between 3600 and 8500 ft in sagebrush, saltbush, conifers, juniper, oak, manzanita, grass, and grease- wood areas in Colorado. Cole (1968:71) shows it across all of extreme southern Utah, arching northward near the borders as it ex- tends into central and southeastern Nevada and central western Colorado. He states that in southern Nevada its nests consist of low gravel mounds or discs, commonly found in hopsage-matrimony vine, shadscale, black- brush, creosote bush, Russian thistle, and other mixed shrubs (1966:5). Ingham (1959, 1963) indicates its habitat in southern Utah as flattened pebble mounds in creosote bush, sagebrush, pinyon-juniper, blackbrush, Joshua trees, bur sage, little rabbitbrush, four-wing saltbush, and shadscale. Allred and Cole (1979:99) found it in southern Utah in several desert shrub types, most commonly in grass communities. There were 160 ants in seven collections taken from low crater mounds not over four inches high and 18 inches to two feet in di- ameter. Eleven ants in one collection were taken from under a log. No typical cleared area occurs around these mounds. Two col- lections were taken in blackbrush, one in as- sociation with grass and Russian thistle. Two collections were in greasewood: one in asso- ciation with Russian thistle and one with grass, herbs, and sagebrush. One collection was taken in herbs, one saltbush, one grass and matchbnish, and one grass near aspen. In 36 recorded Utah habitats two were in mon- tane areas. In 27 recorded elevations between 2500 and 7500 ft, 21 were between 3000 and 5000, two over 7000. These ants are not as aggressive as occidentalis, and some attempt to hide when disturbed. Ants of one colony 502 Great Basin Naturalist Vol. 42, No. 4 found in a habitat of blackbrush, Russian thistle, and grass 35 miles west of U261 along 1LJ95 in San Juan County were harvesting grass seeds on 30 July and carrying them into their nest. The record in northern Utah County col- lected by me is valid. I revisited the exact site a year later, but no ants were present. In the initial collection only 1 1 ants could be found, and they were at 7450 ft under a small aspen limb on an open west-facing, grassy slope be- low an aspen grove. Pogonomyrmex salinus Olsen P. salinus Olsen, 1934, Bull. Mus. Comp. Zool. 77:498; Knowlton 1970:211, 1975:7. Records (Map 24): BEAVER: Desert Range Exp Sta (BY). BOX ELDER: Cedar Crk (City) (KU), Cedar Hill (K75), Curlew Jet, Curlew Valley, Hardup (KU), Kelton (K70), 5 mi SE (KU), 8.3 mi E, 3.3 and 12 mi W, and 39.8, 50.9 and 60.1 mi SW (A), Kelton Pass (K70), Kosmo (K75), Locomotive Spngs (K70), Penrose 7 mi NW, Park Valley (City) 10 mi E (KU), Promontory 20.3 mi W (A), Snowville (K75), Wildcat Hills (US). IRON: Cedar City (RAU). JUAB: Callao 5 and 10 mi E (A), Fish Spngs (KU), Candy 10.1 mi N (A), Topaz Mt (KU), Trout Crk (City) 12.3 mi N (A). MILLARD: Candy 1.7 mi N, and 3.2, 3.5, 8.5, 13.6, 17.2 and 20.1 mi S, Candy rd 0.5 mi N US6-50, Hinckley 14.7 mi W (A). TOOELE: Knolls 3, 3.5, 13, 16.8 and 22.8 mi W, Wendover 5.5 mi E (A). Smith (1979:1355) lists this species from western United States, including Nevada, nesting in a low craterlike mound in pinyon- juniper areas. Cole (1968:107) shows it along the eastern border of Nevada in the northern half of that state but not next to Utah in the southern third of Nevada. He indicates that in southern Nevada it is one of the dominant ants of the pinyon-juniper, replacing occiden- talis at higher elevations, contacting but not overlapping the range of the latter. The nests are saucerlike depressions in cleared areas (1966:6). Knowlton (1975:7) found it associ- ated with sagebrush in northern Utah. There were 510 ants in 20 collections taken from low crater mounds. Sixty-five ants in 4 collections were taken from low, flat mounds, and 40 ants in 2 collections were taken from ground burrows without mounds. Eighty ants in 3 collections were taken from mounds that were larger than the others and typically shaped like those of occidentalis; however, all had openings at the top, only one with a crater. Immatures were found in 5 nests and winged forms in 10. Both imma- tures and males were found together in 4 nests. Fourteen collections were taken in greasewood: 5 in association with halogeton, 2 shadscale, and one halogeton and shadscale. Five collections were in shadscale, 5 shad- scale and halogeton, one halogeton, one halogeton and saltbush, and one pickleweed. Two collections were taken on the shoulder of a road where no vegetation was present. All 44 recorded Utah habitats were in desert areas between elevations of 4222 and 5834 ft. These ants are not aggressive and attempt to hide when disturbed, exhibiting little at- tempt to rescue exposed immature forms. In only two colonies was there indication of ag- gression. In two colonies alongside a paved highway, kernels of wheat were found. This species apparently extends alongside highway US6-50 70 miles eastward from the Nevada border to within about 14 miles of Hinckley in Millard County. At intervals westward from Hinckley the soil is an alka- line clay with greasewood, shadscale, and saltbush. No harvester mounds, even of occi- dentalis, were seen in these areas except along the shoulder of the road where a gravel substrate was present. When the greasewood, saltbush, and alkaline-clay are replaced by desert pavement and shadscale, harvester mounds are present. For 8 miles north of US6-50 alongside the Nevada border toward Candy, only two mounds were seen in the al- kali-clay greasewood habitat up to the shad- scale of the bajada. In the valley east of Kel- ton in Box Elder County in northwestern Utah, where the soil is not as alkaline or clay, harvester mounds are common in greasewood alongside the highway. However, the mounds there are small and frequently covered with greasewood detritus. For many miles south- west of Kelton in the greasewood-shadscale- halogeton habitat, no mounds of salinus were seen except those occasionally in the middle of the hardpacked dirt road. This species with its typical low crater mounds extends along US80 from about 1.5 miles east of Knolls in Tooele County west- ward to Wendover. Typical occidentalis with its higher pyramid mounds extends eastward along 180 from about 3 miles west of Knolls. In this overlap area of about 4.5 miles are some mounds that are intermediate between December 1982 Allred: Ants of Utah 503 the two species types— higher pyramid moimds with top craters. In the area east of the typical crater mounds for a distance of about 1.5 miles, no typical crater mounds were seen but frequent intermediate types were present. The intermediate types were not seen eastward from about 3 miles east of Knolls. Between 40 and 60 miles southwest of Kel- ton in Box Elder County the only mounds found were in the middle of the hard-packed dirt road. Between about 25 and 70 miles west of Hinckley in Millard County along US6-50, wherever mounds of occidentalis were abundant no colonies of salinus were foimd. In a halogeton-shadscale area 12.3 miles north of Trout Creek City in Juab County, colonies of occidentalis were abun- dant in the same area as salinus. In an alkali-greasewood area five miles east of Callao in Juab County a completely flat mound was covered with a dense concen- tration of black chips of desert pavement. In the same habitat five miles farther east a typ- ical mound was covered with black chips of desert pavement, including the inside of the crater. In a shadscale area 14.7 miles west of Hinckley in Millard County a mound was typical occidentalis in shape except that the opening was at the top. However, the mound lacked the crater typical of salinus. These ants demonstrated typical salinus behavior of nonaggressiveness and attempts to hide when disturbed. The mound was conspicuously covered with black chips of gravel that were common in the vicinity as desert pavement. However, the closest source of these to the mound was about 50 ft away. Another mound in the same area was of the same shape but possessed a small crater on the top. This mound was covered by light-colored gravel, probably brought from excavating within the mound. These ants behaved as typical sa- linus. A mound that was an intermediate type— relatively high pyramid with top cra- ter—was found 3.5 miles west of Knolls alongside US80 in Tooele County. The ants behaved as intermediates between salinus and occidentalis—some tried to escape and hide, whereas others crawled around in a manner typical of occidentalis. In series 395, taken 13 miles west of Knolls along US80 in Tooele County, all of the spec- imens examined have typical salinus straight- margin mandibles, but some have the basal part of the scape more typical of occidentalis. Others in the same series have the typical sa- linus scape. In series 397, taken 22.8 miles west of Knolls alongside US80, in Tooele County, two ants have all the teeth rounded and not heavily sclerotized. The distal three teeth are fused together. The basal tooth is not offset and is in straight alignment with the mandibular margin. Two others have the teeth rounded, with the two distal ones fused, but the basal tooth is in straight alignment with the margin of the mandible. In series 405, taken 3.5 miles west of Knolls, the ants have characters typical of both salinus and occidentalis. The basal tooth is consistently in line with the others, not offset. The superior lobe of the scape in many is more rounded (almost like typical salinus) than in occiden- talis, but many specimens have a superior lobe that is more angled, similar to that of oc- cidentalis. In series 510, taken 17.2 miles south of Gandy in Millard County, the ants are typical salinus except that, although some have the superior lobe of the scape rounded as is typical of salinus, others have a lobe that is more angular like occidentalis. In series 548 there are intergrades between salinus and owyheei. One specimen has a sa- linus antenna on one side and an owyheei an- tenna on the other, with a typical salinus petiole. Another has salinus antennae and an owyheei petiole; one has owyheei antennae and salinus petiole. Most others of the series are typical salinus. In series 551, taken 39.8 miles southwest of Kelton, two specimens have owyheei antennae but salinus petioles. In series 546, taken 8.3 miles east of Kelton, some of the ants are typical salinus with ref- erence to the superior lobe of the base of the scape, transverse rugae on the petiole, and the postpetiole. A few specimens have the su- perior lobe of the base of the scape similar to that of owyheei, and on these the transverse rugae on the petiole and postpetiole are not as distinct as on typical salinus, especially on the anterior half of the postpetiole. Pogonomyrmex subnitidus Emery P. occidentalis var. subnitidus Emery, 1895, Zool. Jahrb. Syst. 8:310. i 504 Great Basin Naturalist Vol. 42, No. 4 P. subnitidus: Allred and Cole 1979:99. Records: KANE: Glen Cyn City (AC). SAN JUAN: Mexican Hat 3.2 mi S (A). Smith (1979:1355) lists this species only from Nevada nesting in craterlike mounds, and Cole (1968:114) lists it from western Ne- vada. Allred and Cole (1979:99) found it in southern Utah in ephedra-grass associations. Thirty ants in one collection were taken from a typical occidentalis-type mound in an association of matchbrush and Russian thistle. I was stung on the arm by one of the work- ers, and the reaction was equal to if not more severe than that to occidentalis. Ants of this species are significantly larger than those of occidentalis. The head is much darker than occidentalis, almost black, and the rugae are much closer together, with a little-beaded appearance. The basal tooth on the mandible is almost curved toward the other teeth, and the inner margin of the man- dible is generally straight except that a small hump exists. In series 335, taken 11.6 miles south of Bluff in San Juan County, 24 of the 25 specimens collected have the mandibular teeth worn down, some to the point where no vestiges of teeth are visible. One such specimen has the tips of both mandibles bro- ken off. (1958:167) designates it in Utah as typically a mountain form in dry and rocky situations where sagebrush is present. Ingham (1959) found it in southern Utah in pinyon-juniper and sagebrush in association with Con- oniynna insana. Ten ants in one collection were taken from a burrow imder a log in an association of grass, herbs, aspen, and pine. Ants of Formica fusca were in the same burrow. In 12 re- corded Utah habitats it occurred six times in montane areas. In 8 elevation records be- tween 4535 and 8000 ft it was most abundant under 6000, taken only twice above 6000. Ponera pennsylvanica Buckley P. pennsylvanica Buckley, 1866, Proc. Ent. Soc. Phila. 6:171; Smith 1979:1342. Record: COUNTY UNKNOWN: locality unknown (Sm79). Smith (1979:1342) lists this species from eastern to western United States, including Utah and Colorado, nesting under logs, stones, and other objects. Hunt and Snelling (1975:20) list it from Arizona. Gregg (1963:282) lists it between 3600 and 6970 ft under rocks in pinyon-juniper and cotton- wood-willow habitats in Colorado. Polygerus breviceps Emery P. rufescens breviceps Emery, 1893, Zool. Jahrb. Syst. 7:666; Rees and Grundmann 1940:10; Cole 1942:385; Creighton 1950:558. P. rufescens umbratus: Creighton 1950:560; Grundmann 1958:167; Ingham 1959. Records (Map 25): CACHE: Green Cyn, Logan, Lo- gan Cyn (C42), Providence (KU). GARFIELD: ""Boulder Mt nr Boulder (G58). IRON: Cedar City (159). KANE: Cedar City 24.3 mi E (A). SALT LAKE: Big Cotton- wood Cyn (U). SANPETE: Chester (RG), Majors Flats (KU). SEVIER: Richfield (RG). UINTAH: Bonanza (KU). Smith (1979:1466) lists this species as mid- western to western United States, including Arizona, where it associates with a variety of species of Formica. Gregg (1963:638) lists it from Colorado between 5080 and 8900 ft un- der rocks, logs, and in domes in a variety of habitats. La Rivers (1968:11) lists it from Ne- vada, where Wheeler and Wheeler (1978:396) found it between 6400 and 10,500 ft. Cole (1942:385) indicates its habitat in Utah as under stones, associated with a vari- ety of species of Formica. Grundmann Solenopsis aiirea Wheeler S. geminata var. aurea Wheeler, 1906, Amer. Mus. Nat. Hist., Bull. 22:336. S. aurea: Ingham 1963:86. Record: WASHINGTON: St George (159). Smith (1979:1385) lists this species from southwestern United States, including Ari- zona, in moundless nests in fully exposed situ- ations. Cole (1966:17) found its nests in southern Nevada in open areas in blackbrush and hopsage-matrimony vine habitats. Ing- ham (1963) found it in Joshua trees and four- wing saltbush in southern Utah. Records for Utah range from 2500 to 3000 ft. Solenopsis molesta (Say) Myrmica molesta Say, 1836, Bost. J. Nat. Hist. 1:293. S. molesta: Rees and Grundmann 1940:4; Cole 1942:361; Knowlton 1970:211, 1975:8. S. tnolesta validiuscttla: Cole 1942:361; Grundmann 1958:163; Ingham 1959:58; Beck et al. 1967:72; Knowlton 1970:211, 1975:8. Records (Map 25): BOX ELDER: Cedar Crk (City), Kelton Pass, Snowville (K70), Wildcat Hills (US). December 1982 Allred: Ants of Utah 505 CACHE: Benson (C42), Cornish, Green Cyn (KU), Lo- gan Cyii (C42), Millville, Paradise (KU). CARBON: My- ton rci 15 and 22.7 mi E US6 (A). DAVIS: KavsviUe (C42). DUCHESNE: Currant Crk (C42). EMERY: Greenriver (C42). GARFIELD: Carcass Crk (on Boulder Mt), Boulder (U). GRAND: Dewey (U), Moab (C42). IRON: Cedar Citv 7 mi E (159). JUAB: Topaz Mt (KU). KANE: Kanab (C42), Long Valley (KU), Long Vallev Jet 2 mi W (159), Navajo Wells (BAD). MILLARD: White Valley (C42), Kanosh (US). PIUTE: Fish Lake Jet 1 mi S (U). SALT LAKE: Big Cottonwood Cyn (U), Emigration Cyn, Ft Douglas, Lake Blanche Trail, Little Willow Cyn, Parleys ^Cvn (C42), Salt Lake City (RG). SAN JUAN: Blanding,' Bluff (G58), La Sal Crk (in La Sal Mts) (RG), Mexican Hat (G58). SANPETE: Ephraim Cyn (U). TOOELE: Ibapah (U), losepa, Orrs Ranch, Vernon Crk (C42), Rush Valley (BAD). UINTAH: Drv Fk rd 15 mi N U121 (A), Jensen (BAD), Paradise Park 11 mi S (U). UTAH: Diamond Fk Cyn (KU), Provo (C42), Spanish Fk Cyn, Wanrhodes Cyn (KU). WASATCH: Deer Crk Res, Midway (U), Soldier Summit (BAD) and 7.9 mi N (A). WASHINGTON: Beaver Dam Wash, Diamond Valley (BAD), Grafton (159), Leeds (RG) and 5 mi N, New Har- mony, Zion Nat Park (159). WAYNE: Fruita 5 mi SE (U). WEBER: Ogden (C42). Creighton (1950) lists two races of this spe- cies; the Utah one is likely validiuscula. Smith (1979:1387) lists this species distribu- tion from Canada to eastern and western United States, but does not list an inter- moimtain one. He indicates that it frequently nests in or near nests of other ants, from which it robs food and brood. Hunt and Snelling (1975:22) list it from Arizona. Gregg (1963:373, 375) lists it from Colorado be- tween 3500 and 8378 ft under rocks in a vari- ety of habitats. Cole (1966:17) found its nests in southern Nevada in pinyon-juniper and desert shrub habitats, frequently associated with Pheidole pilifera. He indicates that it nests under stones and logs or under bark, oc- casionally occupying the nests of other spe- cies in Utah (1942:361). In North Dakota Wheeler and Wheeler (1963) found it fre- quently under rocks. In Utah it is found un- der stones in brushy habitats and transition zones, common in desert conditions (Grund- mann 1958:163). Ingham (1959, 1963) found it in southern Utah under stones, logs, and bark in pinyon-juniper, oak, sagebrush, rab- bitbrnsh, squawbush, willow, tamarix, poplar, shadscale, and greasewood. Knowlton (1975:8) found it under stones in grass in northern Utah. There were 247 ants in four collections taken from under rocks. Eggs were found un- der one rock and immature stages under another, both in early July. Ants of Formica haemorrhoidalis were found under the same rock in one instance, and F. fusca and F. pod- zolica in another. In both cases the colonies were separate. Three collections were in sagebrush: one in association with juniper; one grass, legumes, sagebrush, other shrubs, juniper, and pinyon; and one grass, aspen, and pine. One collection was in aspen and fir. In 65 recorded Utah habitats 16 were in montane forest. Of 42 recorded elevations be- tween 2500 and 9000 ft the greatest number, 19, were between 4000 and 5000. Beck et al. (1967:72) found it feeding on dead rodents in seven instances in Utah. Solenopsis salina Wheeler S. salina Wheeler, 1908, Amer. Mus. Nat. Hist., Bull. 24:427; Ingham 1959:59. Records: WASHINGTON: Kolob, Pine Valley (City) (159). Smith (1979:1388) lists this species from western United States, including Colorado, nesting under stones and wood. Hunt and Snelling (1975:22) list it from Arizona. Gregg (1963:378) lists it between 3500 and 7704 ft under rocks and wood in a variety of habitats in Colorado. Cole (1966:17) found it in south- ern Nevada under stones in pinyon-juniper. Ingham (1959) found it in southern Utah un- der stones in oak where it was associated with Formica fusca. Three Utah collections were taken between 6000 and 8000 ft. Solenopsis xyloni McCook S. xyloni McCook, 1879, (In) Comstock, Rpt. Cooton Ins., p. 188; Ingham 1959:58. Records: WASHINGTON: Santa Clara (US), Shivwits Indian Res (159), St George (US). Smith (1979:1389) lists this species from eastern to western United States including Colorado, nesting in earthen mounds or un- der stones and other objects. Hunt and Snell- ing (1975:22) list it from Arizona. Gregg (1963:373) lists it at 4400 ft under rocks in cottonwood-willow and grassy habitats in Colorado. Cole (1966:17) found its nests in southern Nevada at the base of shrubs in creosote bush habitats. Ingham (1959) found it under rocks in creosote bush, poplar, wil- low, datura, and rabbitbrush in southern Utah. Known habitats in Utah are in desert 506 Great Basin Naturalist Vol. 42, No. 4 Stenamma brevicome (Mayr) Aphaenogaster brevicornis Mayr, 1886, Verh. Zool.-Bot. Ges. Wien 36:443. S. brevicome: Cole 1942:363; Knowlton 1975:8. Record: BOX ELDER: Snowville 17 mi SW (KU). CACHE: Logan (C42). Smith (1979:1358) lists this species from eastern to western United States, including Colorado, nesting under stones or wood in wooded areas. Gregg (1963:348) lists it at 3800 ft in deciduous areas in Colorado. Knowlton (1975:8) found it associated with sagebrush and shadscale in northern Utah. Stenamma chiricahua Snelling S. chiricahua Snelling, 1973, Los Angeles Co. Mus., Contr. Sci. 245:7. Record: MORGAN: Morgan (KU). Smith (1979:1358) lists this species from mountains in southern Arizona. Its occur- rence in Utah is questionable. Stenamma diecki Emery S. ivestwoodi diecki Emery, 1895, Zool. Jahrb. Syst. 8:300. S. diecki: Cole 1942:363; Smith 1957:140; Snelling 1973:18; Knowlton 1975:8. Records (Map 25): BOX ELDER: Cedar Hill, Wildcat Hills (K75). CACHE: Blacksmith Fk Cyn, Green Cyn (KU), Logan (C42), Logan Cyn, Trenton (KU). UTAH: American Fk Cyn (U). Smith (1979:1358) lists this species from eastern to western United States, but does not indicate an intermountain state. It nests un- der wood or other objects in wooded areas. Wheeler and Wheeler (1978:391) found it at 6400 ft in Nevada, and frequently under and in wood in North Dakota (1963). Gregg (1963:350) lists it from Colorado between 6050 and 7350 ft under rocks in conifers. In eight recorded Utah habitats it was taken four times in montane forest. Three recorded elevations were between 4460 and 7000 ft. (1975:8) found it associated with sagebrush, rabbitbrush, and junipers in northern Utah. Stenamma impar Forel S. brevicome impar Forel, 1901, Soc. Ent. de Belg., Ann. 45:347. Record: UTAH: Spanish Fk Cyn (KU). Smith (1979:1359) lists this species as east- ern to midwestern United States, and does not indicate an intermountain state. It nests in soil or rotten wood. Stenamma occidentale M.R. Smith S. occidentale Smith, 1957, Amer. Midi. Nat. 57:146; Snelling 1973:25; Smith 1979:1359. Records (Map 25): CACHE: Blacksmith Fk Cyn, Green Cyn, Leeds Cyn, Paradise (KU). UINTAH: Bo- nanza (KU). UTAH: Diamond Fk Cyn (KU). Smith (1979:1359) lists this species from midwest and western United States, including Utah, Colorado, Arizona, and Idaho, nesting under rocks. Gregg (1963:353) lists it from Colorado between 5900 and 8500 ft under rocks in conifers, birch, and oak habitats. Stenamma smithi Cole S. smithi Cole, 1966, Brigham Young U. Sci. Bull. 7(3):7; Knowlton 1975:9; Smith 1979:1359. S. knowltoni: Gregg 1972:35; Knowlton 1975:8. Records (Map 26): BOX ELDER: Cedar Crk Jet (Gr), Cedar Hill (US), Curlew Valley Jet, Hansel Mts, Hard- up, Kelton Pass, Snowville (K75). CACHE: Newton (KU). Smith (1979:1359) lists this species from Utah, Nevada, and Idaho, nesting in sage- brush and juniper duff. Cole (1966:8) found the type series in mixed brush in Nevada. Gregg (1972:38) lists its Colorado habitats as duff of sagebrush, juniper, and greasewood, and in grass. Wheeler and Wheeler (1978:391) found it at 7000 ft in Nevada. Knowlton (1975:8, 9) found it associated with sagebrush, grass, greasewood, and rabbitbrush in northern Utah. Stenamma huachucanum M.R. Smith S. huachucanum Smith, 1957, Amer. Midi. Nat. 57:153; Knowlton 1975:8. Records: BOX ELDER: Hansel Mts, Hardup, Kelton Pass, Snowville (K75). Smith (1979:1359) hsts this species only from Arizona nesting under rocks. Gregg (1963:350) lists it from Colorado at 8300 ft under rocks in mixed forest. Knowlton Tapinoma sessile (Say) Formica sessile Say, 1836, Bost. J. Nat. Hist. 1:287. T. sessile: Rees and Grundmann 1940:6; Cole 1942:372; Hayward 1945:120; Grundmann 1958:165; Ing- ham 1959:68; Beck et al. 1967:73; Knowlton 1970:212, 1975:9. Records (Map 26): BOX ELDER: Bear River City (US), Brigham 0.9 mi E (A), Cedar Crk (City) (K70), Ce- dar Hill (K75), Hansel Mts, Kelton Pass (K70), Locomo- tive Spngs (BAD), Mantua (KU). Raft River S Fk (RG), December 1982 Allred: Ants of Utah 507 Snowville (K75), Taylor Farms (K70), Wildcat Hills (K75). CACHE: Ant Valley, Blacksmith Fk Cyn (KU), Cowley Cyn (C42), Green Cyn, Leeds Cyn (KU), Logan, Logan Cyn (C42), Mendon Cold Spng, Providence, Tony Grove (KU). CARBON: Jet Soldier Summit rd and U33 L7 mi W (A). DAGGETT: Radosovich Ranch (BAD). DAVIS: Antelope Island (US), Farmington (C42). DUCHESNE: Fmitland 5 mi E (US), Johnny Star Flat (BAD), Tabiona n.6 mi E (A). EMERY: Greenriver (US). GARFIELD: Boulder Mt, Henry Mts (G58). IRON: Enoch (BAD), Modena 10.4 mi NE (A), New- castle 1 mi E (159). JUAB: Nephi (US). KANE: Kanab (C42), Long Valley Jet (159). MILLARD: White Valley (C42). MORGAN: Morgan (KU). SALT LAKE: Arthur (US), Big Cottonwood Cyn, Draper (C42). SAN JUAN: Abajo Mts, Blanding, Bluff (G58), Hole-in-the-Rock Cyn (U), Kigalia Ranger Sta (BAD), Monticello 7.6 mi W (A). SEVIER: Monroe (US). SUMMIT: Echo (BAD), Kamas 14.7 mi E, Mirror Lake 17.3 mi N (A). UINTAH: Bo- nanza (KU), Dry Fk rd 13.4 mi N U121, Red Cloud Loop rd 4.2 mi W U44 (A). UTAH: Payson Cyn 12.3 mi up (A), Provo (BAD), Thistle 7.7 mi E (A). WASATCH: Hanna 9.2 mi W, Soldier Summit 3.3 and 10.1 mi N (A), Strawberry Valley (US), Strawberry Res 4 mi S (A). WASHINGTON: Leeds (RG), Pine Valley (City), Santa Clara (159), Toquerville (BAD), Zion Nat Park (159). WAYNE: Hanksville (BAD). Smith (1979:1421) lists this as primarily an eastern species (no intermountain state is listed) that nests mostly under objects in a variety of habitats. Hunt and Snelling (1975:22) list it from Arizona. Gregg (1963:446) lists it from Colorado between 3500 and 10,505 ft under rocks and wood in a variety of habitats, predominantly in conifers. La Rivers (1968:7) lists it from Nevada, where Wheeler and Wheeler (1978:392) found it between 6100 and 10,500 ft. They found it frequently under rocks, also common in wood in North Dakota (1963). Allred and Cole (1971:239) found it in Idaho in associ- ations of wild rye-grass and rabbitbrush-sage- brush-grass-winterfat. Cole (1942:372) in- dicates that in Utah it nests under stones, logs, and bark. Grundmann (1958:165) in- dicates that it is common in Utah in moun- tain brush in large nests usually under stones in shady areas between 4000 and 8000 ft. Ingham (1959, 1963) found it in southern Utah under rocks, logs, and in grass clumps in a variety of vegetative types. Knowlton (1975:9) found it associated with sagebrush, shadscale, greasewood, snowberry, and rab- bitbrush in northern Utah. There were 556 ants in 13 collections taken from under rocks. In one instance Lasius alienus was under the same rock. Eighty ants in 3 collections were taken from under logs. Thirty ants in one collection were taken from a small mound, and 20 in one col- lection singly in an open area. Immatures were found under two rocks in mid-July and under a log in late June. Fourteen collections were taken in sagebrush: 3 in association with grass, 2 snowberry, 2 matchbrush, one grass and herbs, one legumes, and one aspen and conifers. One collection was in a herb meadow with aspen; one grass, herbs, shrubs, and aspen; one herbs, sedges, and conifers; and one grass, herbs, aspen, and conifers. In 70 recorded Utah localities 26 were in mon- tane areas. At 31 recorded elevations be- tween 2500 and 8402 ft, 22 were between 4000 and 6000. Beck et al. (1967:73) found it feeding on dead rodents in 12 instances in Utah. Tetramorium caespitum (Linnaeus) Formica caespitum Linnaeus, 1758, 10th ed. Syst. Nat. 1:581. Record: SALT LAKE: Salt Lake City (KU). Smith (1979:1400) lists this species from eastern to western United States, including Nevada, nesting in a variety of situations. Veromessor lobognathus (Andrews) Messor lobognathus Andrews, 1916, Psyche 23(3):82. V. lobognathus: Wheeler and Wheeler 1967:240; Allred and Cole 1979:99. Records: DUCHESNE: Duchesne 11 mi W (W67). KANE: Glen Cyn City (AC). Smith (1979:1364) lists this species from midwest to western United States, including Colorado and Nevada, nesting under stones. Gregg (1963:354) lists it between 5747 and 6500 ft under rocks in pinyon-juniper and sagebrush areas. Cole (1966:12) found it a common species in pinyon-juniper in south- ern Nevada, nesting under large rocks. Allred and Cole (1979:99, 1971:239) found it in southern Utah and Idaho in various associ- ations of desert shrubs. It was most abundant in rabbitbrush-sagebrush. Wheeler and Wheeler (1963) found it frequently under rocks in North Dakota, and indicate that it is rarely taken, occurs between 7000 ft in the southern part of its range to 2500 ft at the northern part, and is an inhabitant of pinyon- juniper, where it nests under stones (1965:60). They found a nest in Utah imder a mound of 508 Great Basin Naturalist Vol. 42, No. 4 earth at the base of a chimp of grass in an as- sociation of sagebrush, saltbush, and grass (1967:240). Synonymies and Corrections OF Utah Records Names on the left are as written in pub- lished literature or on labels on specimens in collections I examined (many representing heretofore unpublished data), and for pur- poses of this paper are considered to be er- rors in identification unless other authors have treated them as junior synonyms to the names on the right. Acanthomyops claviger = A. interjectus coloradensis = A. interjectus Aphaenogaster subterranea valida = A. occidentalis Brachymyrinex depilis flavescens = B. depilis Camponotus caryae decipiens = C. nearcticiis herculeanus ligniperda noveboracensis = C. novaeboraeensis herculeanus pennsylvanicus = C. herculeanus herculeanus whymperi = C. herculeanus ligniperdus noveboracensis = C. novaeboraeensis maccooki = C. semitestaceus maculatus sansabeanus = C. sansabeanus maculatus sansabeanus torrefactus = C. sansabeanus maculatus vicinus = C. vicinus marginatus nearcticus = C. nearcticus nearcticus decipiens = C. nearcticus pennsylvanicus = C. modoc pennsylvanicus modoc = C. modoc sansabeanus sansabeanus = C. sansabeanus sansabeanus torrefactus = C. sansabeanus sansabeanus vicinus = C. vicinus sansabeanus vicinus luteangulus = C. vicinus sansabeanus vicinus nitidiventris = C. vicinus sylvaticus maccooki = C. semitestaceus Crematogaster coarctata mormonum = C. mormonum lineolata = C. emeryana lineolata cerasi = C. emeryana lineolata coarctata = C. mormonum lineolata coarctata mormonum = C. mormonum lineolata emeryana = C. emeryana lineolata opaca depilis = C. depilis punctulata = C. emeryana vermiculata = C. coarctata Dorymyrmex = Conomyrma bicolor = C. bicolor pvramicus = C. insana pyramicus bicolor = C. bicolor pyramicus flavus = C. insana pyramicus pyramicus = C. insana Eciton = Neivamyrmex sp. = probably N. californicus Formica aliena = Lasius alienus cinerea = F. canadensis cinerea altipetens = F. altipetens cinerea canadensis = F. canadensis cinerea cinerea altipetens = F. altipetens cinerea lepida = F. canadensis cinerea neocinerea = F. canadensis claviger = Acanthom.yops claviger crinata = F. criniventris crinoventris = F. criniventris flava = Lasius nearcticus foreliana = F. gnava fusca argentata = F. argentea fusca argentea = F. argentea hisca cinerea = F. altipetens fusca densiventris = F. densiventris fusca gelida = F. neorufibarbis fusca neoclara = F. neoclara fusca neorufibarbis = F. neorufibarbis fusca pruinosa = F. neoclara fusca subaenescens = F. fusca fusca subpolita = F. subpolita fusca subpolita neogagates = F. neogagates fusca subpolita perpilosa = F. perpilosa fusca subsericea = F. fusca herculeana = Camponotus herculeanus insana = Conomyrma insana integra = F. haemorrhoidalis integra haemorrhoidalis = F. haemorrhoidalis integroides coloradensis = F. integroides integroides propinqua = F. integroides integroides planipilis = F. integroides laevigatus = Camponotus laevigatus latipes = Acanthomyops latipes marcida = F. fusca microgyna = F. rasilis microgyna rasilis = F. rasilis moki = F. xerophila moki grundmanni = F. xerophila moki xerophila = F. xerophila montana = F. canadensis neogagates lasioides vetula = F. lasioides neorufibarbis gelida = F. neorufibarbis niger = Lasius niger novoboracensis = Camponotus novaeboraeensis obscuriventris clivia = F. obscuriventris oreas comptula = F. oreas oreas oreas = F. oreas pallidefidva nitidiventris = F. pallidefidva pallidefulva nitiventris = F. pallidefulva pennsylvanica = Camponotus modoc planipilis = F. integroides propinqua = F. integroides pruinosa = F. neoclara rasilis spicata = F. densiventris rufa aggerans = F. obscuripes rufa clivia = F. obscuriventris rufa coloradensis = F. integroides rufa haemorrhoidalis = F. haemorrhoidalis rufa laeviceps = F. laeviceps rufa melanotica = F. obscuripes rufa muscescens = F. mucescens rufa obscuripes = F. obscuripes rufa obscuriventris integroides = F. integroides December 1982 Allred: Ants of Utah 509 nifibarbis gnava = F. gnava nifibarbis occidua = F. occidua sangiiinea = F. subnuda sanguinea obtusopilosa = F. obtusopilosa sangiiinea pubenila = F. puberula sanguinea nibicunda subnuda = F. subnuda sanguinea subnuda = F. subnuda sansabeanus = Caniponotus sansabeanus sessilis = Tapinoma sessile subaenescens = F. subnitens subpolita camponoticeps = F. subpolita subpolita ficticia = F. subpolita subsericea = F. fusca truncicola integroides = F. integroides truncicola integroides coloradensis = F. integroides truncicola integroides haemorrhoidalis = F. haemorrhoidalis truncicola niucescens = F. mucescens truncicola obscuriventris = F. obscuriventris tmncicola obscuriventris aggerans = F. obscuriventris umbrata = Lasius umbratus vinculans = F. neogagates whyniperi alpina = F. whymperi Iridomyrniex analis = I. pruinosus pminosus analis = I. pruinosus pruinosus pruinosus = I. pruinosus pminosus testaceus = I. pruinosus Lasius alienus americanus = L. alienus americanus = L. niger americanus sitkaensis = L. pallitarsus claviger = Acanthomyops coloradensis flavus = L. nearcticus flavus claripennis = L. nearcticus flavus microps = L. nearcticus flavus nearcticus = L. nearcticus interjectus = Acanthomyops interjectus latipes = Acanthomyops latipes murphyi = Acanthomyops murphyi neoniger = L. niger niger alienus americanus = L. alienus niger americanus = L. alienus niger neoniger = L. niger niger sitkaensis = L. pallitarsus pilosus = L. vestitus sitkaensis = L. pallitarsus umbratus aphidicola = L. umbratus umbratus mixtus aphidicola = L. umbratus umbratus subumbratus = L. subumbratus Leptothorax acervorum canadensis = L. muscorum acervorum canadensis yankee = L. muscorum acervorum crassipilis = L. crassipilis canadensis = L. muscorum canadensis yankee = L. muscorum nevadensis nevadensis = L. nevadensis pilifera = Pheidole pilifera rugatulus brunnescens = L. rugatulus rugatulus rugatulus = L. rugatulus sitkaensis = Lasius pallitarsus tricarinatus tricarinatus = L. tricarinatus tricarinatus neomexicanus = L. tricarinatus Liometopum apiculatum luctuosum = L. occidentale luctuosum = L. occidentale microcephalum occidentale = L. occidentale occidentale luctuosum = L. occidentale tricarinatus = L. occidentale Myrmecocystus melliger = M. mendax melliger mendax = M. mendax melliger orbiceps = M. mendax melliger semiruhis = M. kennedyi mexicanus hortideorum = M. mexicanus mexicanus mojave = M. testaceus mexicanus navajo = M. navajo mojave = M. testaceous Myrmica brevinodis = M. incompleta brevinodes brevispinosa = M. brevispinosa brevinodes discontinua = M. brevispinosa brevinodis sulcinodoides = M. incompleta californica = Pogonomyrmex californicus emeryana emeryana = M. emeryana emeryana tahoensis = M. emeryana fracticornis = M. lobicornis hamulata hamulata = M. hamulata hunteri = Manica hunteri incompleta incompleta = M. incompleta lineolata = Crematogaster emeryana lobicornis fracticornis = M. lobicornis lobicornis lobifrons = M. lobicornis lobifrons = M. lobicornis mojave = M. testaceus molesta = Solenopsis molesta mutica = Manica mutica occidentalis = Pogonomyrmex occidentalis rubra brevinodis = M. incompleta rubra brevinodis brevispinosa = M. brevispinosa rubra brevinodis sulcinodoides = M. incompleta rubra scabrinodis fracticornis = M. lobicornis sabuleti americana = M. americana sabuleti hamulata = M. hamulata scabrinodis = M. monticola scabrinodis brevinodis = M. emeryana scabrinodis lobocornis fracticornis = M. lobicornis scabrinodis mexicana = M. emeryana scabrinodis schenki emeryana = M. emeryana scabrinodis schenki monticola = M. monticola scabrinodis sulcinodoides = M. emeryana schenecki emeryana = M. emeryana Novomessor albisetosus = Aphaenogaster uinta Pheidole bicarinata buccalis = P. bicarinata bicarinata longula = P. bicarinata bicarinata vinelandica = P. bicarinata bicarinata paiute = P. bicarinata californica oregonica = P. californica longula = P. bicarinata morrisi dentata = P. dentata pilifera artemisia = P. pilifera pilifera coloradensis = P. pilifera pilifera pilifera = P. pilifera sitarches sitarches = P. sitarches sitarches soritis = P. sitarches soritis = P. sitarches 510 Great Basin Naturalist Vol. 42, No. 4 Pogonomyrmex barbatus fuscatus = P. barbatus barbatus marfensis = P. barbatus barbatus molefaciens = P. barbatus barbatus rugosus = P. rugosus californicus maricopa = P. maricopa occidentalis comanche = P. occidentalis occidentalis occidentalis = P. occidentalis occidentalis owyheei = P. owyheei occidentalis subnitidus = P. subnitidus occidentalis utahensis = P. occidentalis Polygerus rufescens = P. breviceps ru.'escens breviceps = P. breviceps rufescens fusca subumbratus = P. breviceps rufescens umbratus = P. breviceps Ponera coarctata pennsylvanica = P. pennsylvanica opaciceps = Hypoponera opaciceps trigona = P. pennsylvanica trigona opacior = Hypoponera opacior Solenopsis molesta validiuscula = S. niolesta Stenamma knowltoni = S. sniithi Symmyrmica chamberlini = Formicoxenus chamberlini Tapinoma pruinosum = Iridomyrmex pniinosus Literature Cited Allred, D. 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Additions, corrections, and new synonymy. J. Tennessee Acad. Sci. 30(l):49-50. 1956. Observations of some members of the genus Pheidole in the southwestern United States with synonymy. J. Tennessee Acad. Sci. 31(2): 112-18. 1957. Notes on western ants. J. New York Ent. Soc. 65:129-31. 1966. Ants of the Nevada Test Site. BYU Sci. Bull. Biol. Ser. 7(3): 1-27. 1968. Pogonomyrmex harvester ants: a study of the genus in North America. Univ. of Tennessee Press, Knoxville, 222 pp. Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool. 104:1-585. Cresson, E. T. 1874. Report on collections of Hymenop- tera made in portions of Nevada, Utah, Colorado, New Mexico, and Arizona during the years 1872-1874. Geogr. Explor. West of 100th Me- ridian, Washington, D.C., 1875:705-36. Francoeur, a. 1973. Revision taxonomique des especes Nearctiques du Groupe fusca, genre Formica (Formicidae, Hymenoptera). Mem. Soc. Entomol. Quebec, Mem. 3, 316 pp. 1974. Notes for a revision of the ant genus For- mica. 1. New identifications and synonymies for some Nearctic specimens from Emery, Forel and Mayr collections. Ent. News 85(9 & 10):257-64. Garrett, A. O. 1910. Honey ants in Utah. Science 32:342-3. Gregg, R. E. 1963. The ants of Colorado. Univ. Colo- rado Press, Boulder, Colo. 792 pp. 1972. A new species of Stenamma from Utah. Great Basin Nat. 32(l):35-9. Grundmann, A. W. 1939. The ants of Salt Lake County (with a list of the known ants of Utah). Unpub- lished thesis. Univ. of Utah. 1952. A new Brachymyrmex from northern Utah. J. Kansas Ent. Soc. 25:117. 1958. Ants. Pages 161-167 in Preliminary report on biological resources of the Glen Canyon Res- ervoir. Univ. Utah Anthr. Papers, No. 31. Hayward, C. L. 1945. Biotic communities of the south- ern Wasatch and Uinta mountains, Utah. Great Basin Nat. 6(1-4):1-124. Hunt, J. H., and R. R. Snelling. 1975. A checklist of the ants of Arizona. J. Arizona Acad. Sci. 10(l):20-3. Ingham, C. D. 1959. Ants of the Virgin River Basin, southwestern Utah. Unpublished thesis. Univ. of Utah. 140 pp. 1963. An ecological and taxonomic study of the ants of the Great Basin and Mohave Desert re- gions of southwestern Utah. Unpublished dis- sertation. Univ. of Utah. 212 pp. James, F. K. 1950. The ants of Red Butte Canyon. Un- published thesis. Univ. of Utah. Knowlton, G. F. 1946. Birds feeding on ants in Utah. J. Econ. Ent. 39:547-8. 1963. Western harvester ant control for Utah. Utah State Univ. Ext. Leafl. 48(Rev.), p. 1. 1970. Ants of Curlew Valley. Proc. Utah Acad. Sci., Arts and Letters 47(1):208-12. 1975. Ants of Curlew Valley, Utah and Idaho. Utah State Univ. Ecol. Ctr., Terr. Arth. Ser. 13, 10 pp. Knowlton, G. F., and W. P. Nye. 1946. Lizards feeding on ants in Utah. J. Econ. Ent. 39(4):546-7. La Rivers, I. 1968. A first listing of the ants of Nevada. Biol. Soc. Nevada, Occ. Papers, 12 pp. Olsen, O. W. 1934. Notes on the North American har- vesting ants of the genus Pogonomyrmex Mayr. Bull. Mus. Comp. Zool. 77(8):493-514. December 1982 Allred: Ants of Utah 511 Rees, D. M., and a. W. Grundmann. 1940. A prelimi- nary list of the ants of Utah. Bull. Univ. Utah 31(5): 1-12. Smith, D. R. 1979. Family Formicidae. Pages 1323-1467 in Catalogue of Hymenoptera in America North of Mexico. Smithsonian Inst. Press, Wash., D.C. Smith, M. R. 1952. North American Leptothorax of the tricarinatus-texanus complex. J. New York Ent. Soc. 50:96-106. 1953. A new Pheidole (subgenus Ceratopheidole) from Utah. J. New York Ent. Soc. 61:143-6. 1957. Revision of the genus Stenamma Westwood in America north of Mexico. Amer. Midi. Nat. 57:133-74. Snelling, R. R. 1970. Studies on California ants, 5. Revi- sionary notes on some species of Camponotus, subgenus Tanaernymiex. Proc. Ent. Soc. Wash- ington 72(3):390-7. 1973-a. The ant genus Conomynna in the United States. Los Angeles County Nat. Hist. Mus., Contr. 238, 6 pp. 1973-b. Studies on California ants, 7. The genus Stenamma. Los Angeles County Nat. Hist. Mus., Contr. 245, 38 pp. 1976. A revision of the honey ants, genus Myr- mecocystus. Los Angeles County Mus. Nat. Hist., Sci. Bull. 24, 163 pp. Weber, N. A. 1947. A revision of the North American ants of the genus Myrmica. Ann. Ent. Soc. Amer. 40:437-47. Wheeler, G. C, and J. Wheeler. 1963. The ants of North Dakota. Univ. of North Dakota Press. Grand Forks. 326 pp. 1965. Veromessor lohognathus: third note. J. Kan- sas Ent. Soc. 38(1):55-61. 1967. Veromessor lohognathus: fourth note. J. Kansas Ent. Soc. 40(2):238-41. 1970. The natural history of Manica. J. Kansas Ent. Soc. 43(2): 129-62. 1977. North Dakota ants updated. Desert Res. Inst. Publ., Reno, Nevada. 27 pp. 1978. Mountain ants of Nevada. Great Basin Nat. 38(4):379-96. Wheeler, W. M. 1904. Three new genera of inquiline ants from Utah and Colorado. Bull. Amer. Mus. Nat. Hist. 20:1-17. Wilson, E. O. 1955. A monographic revision of the ant genus Lasius. Bull. Mus. Comp. Zool. 113(1): 1-205. Wing, M. W. 1968. Taxonomic revision of the nearctic genus Acanthomyops. Cornell Univ. Agric. Expt. Sta., Mem. 405. 173 pp. Woodbury, A. M. 1952. Gazetteer of Utah localities and altitudes. Div. Biol., Univ. of Utah. Salt Lake City. 216 pp. VEGETAL RESPONSES AND BIG GAME VALUES AFTER THINNING REGENERATING LODGEPOLE PINE' D. D. Austin- and Philip J. Urness- Abstract.— Understory vegetal response was found to significantly increase with the degree of thinning in an early regenerating, dense stand of lodgepole pine (Piniis contorta). The value of the increased vegetation for deer and elk was determined to be important through comparisons with known dietary and habitat preferences. Following removal of mature lodgepole pine stands, regeneration is frequently dense and results in early stagnation (Forest Service 1962). Although thinning of young stands of- ten increases the rate of growth (Trappe 1959) and harvested yields (Wikstrom and Wellner 1961), other values should be consid- ered, especially when cost /benefit ratios for timber are marginal. Increases in forage pro- duction are a potential additional value; however, the relationship between forest thinning and big game habitat values remains ambiguous (Wallmo and Schoen 1981:445). This paper reports the response of understory vegetation four years following thinning treatments in a dense, 16-year-old lodgepole pine stand and, using previously determined diets and habitat preferences, assesses the po- tential value for deer and elk. Area and Methods The study was on the Ashley National For- est in northeastern Utah near East Park Res- ervoir at 2700 m elevation. Lodgepole pine covers 92 percent of the area, which is an un- dulating upland draining to the south. Natural regeneration of forest stands fol- lowing harvest or fire in the area has resulted in dense stands of trees, usually requiring thinning to prevent stand stagnation. This study was conducted on one such stand bull- dozed and broadcast burned for wildlife and timber values in 1960 and 1961. The regen- erated stand in 1976 had a density of 6200 stems per ha and mean height of 2 m com- pared to the adjacent untreated stand with about 8500 stems per ha and 10 m height. Three replicates of four macroplots, each 20 X 20 m with 4 m buffer strips, were ar- ranged in a randomized block design. Clear- cut, heavy thinning, moderate thinning, and control treatments were established during August 1976 (Fig. 1); trees were handcut and removed. The heavy and moderate thinning treatments left about 1100 and 2200 stems per ha, respectively, which compare with about 1300 stems per ha in nearby stands scheduled for logging and 2000 stems per ha for estimated maximum yield in the Rocky Mountains (Forest Service 1962). Vegetal production and ground cover were deter- mined during August 1976 and 1980 using the microplot-macroplot approach (Poulton and Tisdale 1960) with the modifications of Deschamp et al. (1979). Estimates were re- corded on each of 40 microplots (20 X 50 cm) within each macroplot, with every tenth plot subsequently clipped and weighed for double sampling regression analysis. The 1980 data were subjected to a covariance analysis using 1976 as the covariant. Vegetal Change The response of understory vegetation to tree removal was determined using four in- dices (Table 1). The first two indices, produc- tion and ground cover, indicate the amount of vegetal change, and the latter two, density 'This study was partially supported by funds from the Pittman-Robertson Act under Utah Division of Wildlife Resources Project W-105-R. 'Department of Range Science, Utah State University, Logan, Utah 84322. 512 December 1982 Austin, Urness: Vegetal Responses 513 %.\ )^.-^^ 514 Great Basin Naturalist Vol. 42, No. 4 and number of species, reflect community complexity. With the four indices the mean response between 1976 and 1980 was gener- ally greatest in the clearcut, followed by the heavy thinning, moderate thinning, and control. Mean understory production increased 82 percent on the clearcut, 8 percent on the heavy thinning, 2 percent on the moderate thinning, and the control decreased 18 per- cent. Ground cover increased a mean 102 percent on the clearcut, 47 percent on the heavy thinning, 17 percent on the moderate thinning, and 14 percent on the control. The differences in production and cover due to treatment effects were significant (p<.05). Species density, the mean number of spe- cies encountered on the 0.1 m^ plots, and the number of species present per macroplot showed similar trends. Density increased 59, 43, 26, and 16 percent on the clearcut, heavy thinning, moderate thinning, and control, re- spectively, and the mean number of species increased 5.4 on the clearcut and heavy thin- ning, 4.0 on the moderate thinning, and 1.6 on the control. However, responses due to treatment effects were not significant, al- though species density approached signifi- cance (P<. 06). Means adjusted for pretreatment condition showed significant differences between treat- ments (Table 1). The adjusted means repre- sent the expected values in the fourth year following treatments that would have re- sulted had initial data on all macroplots with- in replications been equal. The clearcut treatment showed significant increases (P<.05) in indices' values over those of the control in production, cover, and density. with number of species approaching signifi- cance (P<.08). Similarly, the heavy treat- ment was significant in cover, density, and number of species, and production ap- proached significance (P<.07). None of the values in the moderate thinning were signifi- cant (P<.05); production became significant at P<.10 and number of species at P<.09, however. These results show a substantial increase in the amount of understory forage and plant community complexity following clearcut and heavy thinning treatments. Furthermore, although the control showed a decrease in production and only a slight increase in cov- er, density, and number of species, the clear- cut and heavy thinning treatments, in con- trast, showed a positive change in production and much larger increases in the other in- dices. The moderate treatment had lesser in- creases. Although each of the four indices evaluated in this study is useful in describing community composition, it is apparent that production and cover are more sensitive in detecting changes. Similar results have been reported by Basile and Jensen (1971) and Regelin et al. (1974) in clearcut areas of lodgepole pine forests elsewhere. Value to Big Game Although a treatment may result in signifi- cant increases in plant production, unless the increase is within preferred grazing areas and composed of species palatable to potential grazers, changes in forage production are in- consequential. Of the five major vegetal seg- ments within the study area— wet and dry meadow and mature, stagnated, and regen- Table 1. Mean indices of understory vegetal changes: production (kg/ha), ground cover (%), species density (spe- cies/0.1 m^), and number of species (species/macroplot). Production Cover Density Number Treatment 1976-1980 1976-1980 1976-1980 1976-1980 Clearcut Heavy thinning Moderate thinning Control Clearcut Heavy thinning Moderate thinning Control 181-329 195-210 172-176 180-147 330* 196^' 1871' 149b 21-43 20-30 20-24 19-21 4P 3* 23bc 23^ 2.8-4.4 2.5-3.6 2.7-3.3 2.3-2.7 4.2^ 3.T''' 3.3bc 2.9^ 13-19 11-17 9-13 10-11 le^'^ le^b ly'^ 13^ a,b,C£)iffgfgp( letters are Adjusted Means and indicate significance at p<.05 within columns. December 1982 Austin, Urness: Vegetal Responses 515 erating lodgepole pine forest— regenerating lodgepole pine was the most preferred habi- tat for deer (Deschamp et al. 1979), and it was second only to the wet meadow for elk (Collins et al. 1978). Thus, increases in pro- duction due to thinning would occur in habi- tats favored by big game. Potential forage benefits were assessed by comparing the 1976 and 1980 production data of major species (Table 2) with the cor- responding dietary preferences for deer (Deschamp 1977) and elk (Collins 1977). For- age preferences were obtained from the ratio of percent diet composition to percent avail- able production (Neff 1974); and preference categories (Table 2), although arbitrarily de- termined, corresponded to animal selection of forage species under free-ranging field conditions. In response to thinning, most forb species increased in production or remained about the same. Production of grass and sedge species also increased in production ex- cept short-stemmed sedge {Carex brevipes), which decreased. Conversely, production of browse species tended to show little response to treatment. Generally deer and elk showed a prefer- ence for most browse and forb species, and grasses and sedges were rejected (Table 2). Consequently, the increased production of grasses and sedges would have little benefit to big game. Production increases in forbs could be highly beneficial, particularly since forbs comprised the large majority of the diets; deer 94 percent (Deschamp et al. 1979) and elk 86 percent (Collins et al. 1978). The small response of browse species would not likely affect the diet. In densely forested areas where natural openings are few, created openings become important as foraging sites (Wallmo et al. 1972, Regelin et al. 1974, Hershey and Leege 1976). However, as regeneration begins to dominate site productivity, understory vege- tation declines (Basile 1975). Maximum un- derstory production in lodgepole pine forests occurred only 10-11 years following either timber harvest (Basile and Jensen 1971) or fire (Lyon 1976) disturbance. Consequently, thinning treatments would lengthen the ef- fective forage-producing interval in forest succession. Table 2. Major plant species within treatment areas, initial production (kg/ha), production after four grazing sea- sons, and deer and elk diet preferences. Species Control Moderate Heavy Clearcut Preference' 19761980 1976 1980 1976 1980 1976 1980 Deer Elk Forbs Antennaria spp. Arnica cordifolia Aster chilensis Astragalus decitmbens Stellaria jamesiana Taraxacum officinale 12 others Total Grasses and Sedges Carex brevipes Carex geyeri Poa spp. Sitanion hystrix 5 others Total Browse Populus tremuloides Rosa nutkana Salix spp. 4 others Total Total 0.1 1.1 23.8 0.0 0.0 14.4 3.8 0.0 0.0 6.1 4.1 5.2 0.3 7.0 2.8 7.0 3.7 7.2 18.5 -l- 1.1 15.2 0.6 5.9 -I- -I- 0.7 1.5 2.7 4.2 4.8 3.6 8.4 16.3 -1- + -1- 38.4 39.7 19.9 33.5 20.1 28.2 26.5 ,37.9 0 — 6.3 9.1 7.9 12.4 6.7 9.9 6.5 8.8 + + 18.1 6.5 18.1 14.9 34.6 25.2 15.8 38.1 -1- + 17.7 29.1 13.9 27.0 6.1 17.0 8.9 18.1 -1- -1- 89.4 97.2 66.9 101.8 80.4 102.8 73.9 143.6 47.4 23.2 60.5 19.2 62.6 41.8 56.8 42.0 2.8 12.2 0.0 0.9 11.7 7.6 1.9 0.0 72.3 .39.1 81.7 6.0 4.9 0.0 0.0 18.4 4.7 0.0 0.0 18.2 10.9 23.1 21.3 17.0 0.0 1.1 58.6 6.4 8.4 0.0 0.6 15.4 3.7 14.6 0.0 0.0 80.9 6.9 25.5 1.1 0.3 7.7 2.1 4.4 24.9 10.2 40.7 0.2 3.3 47.7 5.8 0.0 13.9 80.4 72.4 148.7 11.9 14.2 0.3 0.2 ,33.8 26.6 0.0 21.9 2.9 10.2 ,35.0 0.9 23.9 1.6 10.5 ,36.9 179.9 147.2 171.7 175.8 195.1 209.8 181.3 .329.2 + + + + + 'Ratio of % Diet-% production; 0 = no preference (.75-1.50), + = preferred (1.51-4.00), + + = highly preferred (>4.00), - = rejected (.50-.74), — = highly rejected (<.50). 516 Great Basin Naturalist Vol. 42, No. 4 Our findings indicated an inverse relation- ship between stand density following thin- ning and understory vegetal production. Al- though complete tree removal is untenable on tracts of high site quality, in areas of low timber potential, particularly in stagnated stands, permanent, small openings, consistent with scenic, wildlife, and watershed values (Wyoming Forest Study Team 1971), may be justifiably incorporated into the management plan. Furthermore, both heavy and moderate thinning of regenerating lodgepole pine stands must be considered practical treat- ments for maintaining or slightly increasing the amount as well as the longevity of the forage resource, particularly when contrasted to the control, which showed a decline in for- age production. Literature Cited Basile, J. V. 1975. Forage productivity in the lodgepole pine type. Pages 246-263 in D. M. Baunigartner, ed., Management of Lodgepole Pine Ecosystems Symposium proceedings. Washington State Univ. Coop. Ext. Serv. 825 pp. Basile, J. V., and C. E. Jensen. 1971. Grazing potential on lodgepole pine clearcuts in Montana. USDA For. Serv. Res. Pap. INT-98. 11 pp. Collins, W. B. 1977. Diet composition and activities of elk on different habitat segments in the lodgepole pine tvpe, Uinta Mountains, Utah. Ut. Div. Wildl. Res. Pub. No. 77-18. 74 pp. Collins, W. B., P. J. Urness, and D. D. Austin. 1978. Elk diets and activities on different lodgepole pine habitat segments. J. Wildl. Manage. 42:799-810. Deschamp, J. A. 1977. Forage preferences of mule deer in the lodgepole pine ecosystem, Ashley National Forest, Utah. Utah Div. Wildl. Res. Pub. No. 77- 16. 75 pp. Deschamp, J. A., P. J. Urness, and D. D. Austin. 1979. Summer diets of mule deer from lodgepole pine habitats. J. Wildl. Manage. 43:154-161. Forest Service. 1962. Silvics of forest trees of the United States. Pages 373-387 in USDA Agr. Handbook 271. 762 pp. Hershey, T. J., AND T. A. Leege. 1976. Influences of log- ging on elk on summer range in north central Idaho. Pages 7.3-80 in J. M. Peek and S. R. Hieb, eds., Elk-Logging-Roads Symposium. Univ. of Idaho, Fish, Wildl. and Range Exp. Sta. 1.34 pp. Lyon, L. J. 1976. Vegetal development on the Sleeping Child burn in western Montana, 1961 to 1973. USDA For. Serv. Res. Pap. INT-184. 24 pp. Neff, D. J. 1974. Forage preferences of trained mule deer on the Beaver Creek watersheds. Arizona Game & Fish Dept. Spec. Rep. 4. 61 pp. Poulton, C. E., and E. W. Tisdale. 1961. A quan- titative method for the description and classifica- tion of range vegetation. J. Range Manage. 14:13-21. Regelin, W. L., O. C. Wallmo, J. Nagy, and D. R. DiETZ. 1974. Effect of logging on forage values for deer in Colorado. J. For. 72:282-285. Trappe, J. M. 1959. Lodgepole pine clearcuts in north- eastern Oregon. J. For. 57:420-423. Wallmo, O. C, W. L. Regelin, and D. W. Reichart. 1972. Forage use by mule deer relative to logging in Colorado. J. Wildl. Manage. 36:1025-1033. Wallmo, O. C, and J. W. Schoen. 1981. Chapter 11, Part 2. Forest Management for deer. Pages 434-448 in Mule and black-tailed deer of North America. Wildl. Manage. Institute, Univ. of Ne- braska Press, Lincoln. 605 pp. WiKSTROM, J. H., AND C. A. Wellner. 1961. The op- portunity to thin and prune in the northern Rocky Mountain and intermountain regions. USDA For. Serv. Res. Pap: INT-61. 14 pp. Wyoming Forest Study Team. 1971. Forest manage- ment in Wyoming— timber harvest and the envi- ronment on the Teton, Bridger, Shoshone, and Bighorn national forests. USDA For. Serv., Og- den. 41 pp. HABITAT MANIPULATION FOR REESTABLISHMENT OF UTAH PRAIRIE DOGS IN CAPITOL REEF NATIONAL PARK' Rodney L. Player-" and Philip J. Urness' Abstract.— Utah prairie dogs were transplanted onto the site of a former colony, located in Capitol Reef National Park, Utah. Shrubs on the site were significantly taller than those found on active colonies in similar habitat located on the Awapa Plateau. Therefore, the transplant site afforded a test of the hypothesis that shrub height is a major inhibitory factor affecting occupation of sites by prairie dogs. Four sites of 5 ha each were used. Vegetation treatments— rot obeating, railing, and 2,4-D herbicide— were carried out on three of the sites and the fourth was used as a control. Shrub height and percent cover were significantly reduced on all three treatment sites. Posttreatment effects on the vegetation showed that the greatest percent moisture of the herbage was found on the railed site, fol- lowed by the herbicide, rotobeaten, and control sites. Measurements of the visual obstructions to prairie dogs showed that the rotobeaten site had the greatest visibility, followed by the railed, herbicide, and control sites. Prior to release of prairie dogs on the study area, 200 artificial burrows per treatment were dug, using a power au- ger. In early summer, 1979, 200 Utah prairie dogs were live-trapped near Loa, Utah. An equal number by sex and age class were released on each treatment. In 1979 a significantly higher number of animals reestablished on the rotobeaten site. In 1980 and 1981 the rotobeaten and railed sites had significantly higher prairie dog numbers than the other sites. Reproduction occurred on both the rotobeaten and railed sites in 1980 and 1981. Results indicated that, when transplanting animals onto sites of former colonies presently overgrown with shrubs, the chances of a suc- cessful transplant could be increased by first reducing shrub height and density. The Utah prairie dog {Cynomys parvidens), endemic only to Utah, is presently found in six counties in the south central part of the state (Elmore and Workman 1976). Since 1920 the area occupied by the Utah prairie dog has declined by an estimated 87 percent and their numbers have also declined from an estimated 95,000 in 1920 to an actual count of 3,429 in 1976 (Collier and Spillett 1973). As a result of this decline, the Utah prairie dog was classified as an endangered species in 1968, delisted in 1972, and subsequently reinstated in 1973 (Bureau of Sport Fisheries and Wildlife 1968, 1972, 1973). Possible reasons for the decline in popu- lation and the reduction in range of the Utah prairie dog, as listed by Collier and Spillett (1972), are: purposeful poisoning, disease, drought, shooting, predation, and habitat changes. Poisoning is thought to be the most important factor that has influenced the dis- tribution and abundance of the Utah prairie dog in the past 45 years. Toxicants have been used to eliminate the species from approx- imately 8000 hectares (Collier and Spillett 1972). Population reductions corresponding to periods of intensive poisoning have oc- curred in 1933, 1950, and 1960. However, federal agencies have not used toxicants to control Utah prairie dogs since 1963 (Collier and Spillett 1973). Because of its classifica- tion as an endangered species, the use of tox- icants for population control has been pro- hibited since 1968. Prairie dogs of all species are restricted to habitat of relatively open plant communities with short-stature vegetation (Allan and Osborn 1949, Koford 1958, Fitzgerald and Lechleitner 1974, Collier 1974, Crocker- Bedford and Spillet 1977). According to Col- lier (1974), Utah prairie dogs prefer areas with vegetal cover shorter than 31 cm. Ap- parently this is due to the fact that prairie dogs are dependent upon visual surveillance of their environment to guard against pred- ators and for intraspecific interactions (Fitz- gerald and Lechleitner 1974). Prairie dogs have extended their range into areas where 'This project was funded by the National Park Service, Cooperative Research Unit, College of Natural Resources, Utah State Universit)'. The authors thank the Utah Division of Wildlife Resources, Nongame Wildlife Program, for their cooperation. 'Range Science Department, Utah State University. Logan. Utah 84322. 'Present address: Forest Service, USD.^, .\shlev National Forest. Duchesne, Utah 84021. 517 518 Great Basin Naturalist Vol. 42, No. 4 i the tall, dense, native vegetation has been re- duced by domestic animals and agriculture (Schaffner 1929, Osborn 1942). The converse of this has also been known to occur. A colo- ny of prairie dogs was eliminated when tall, dense vegetation encroached a site after grazing was stopped (Allan and Osborn 1949). The recent elimination of the Utah prairie dog in the Escalante Desert was at least in part attributed to an invasion of woody spe- cies (Collier and Spillett 1973). Snell and Hlavachick (1980) reported that a colony of black-tailed prairie dogs (C. ludovicianus) was reduced in size from 110 acres to 12 acres by allowing cattle to heavily graze the pasture containing the colony in the early spring (thus competing with the prairie dogs for forage) and resting the pasture during June, July, and August, allowing the warm season plants to grow rapidly, creating a visual barrier to the prairie dogs. A general decrease in grasses and an in- crease in brushy species has been observed in the Great Basin since settlement in the mid- 1800s (Pickford 1932, Cottam and Evans 1945, Blaisdell 1953, Ellison 1960, Tueller and Blackburn 1974). Furthermore, the major foods of prairie dogs (herbaceous species) tend to decline in association with highly competitive, xerophytic shrubs such as big sagebrush {Artemisia tridentata), rabbitbrush (Chrysothamnus spp.), and various other shrubs (Ellison 1960, Collier and Spillett 1973, Tueller and Blackburn 1974). This is a result of grazing practices and fire suppres- sion (Pickford 1932, Smith 1949). It should be noted that vegetational changes could have occurred on sites of both occupied and unoc- cupied colonies. Therefore, although the veg- etation on colonies that were eliminated by poisons, disease, predation, shooting, or drought was conducive to prairie dog exis- tence at the time of extirpation, it is possible that subsequent vegetational changes have taken place such that the site is no longer suitable for reestablishment of the colony. Of the six factors affecting populations of Utah prairie dogs, two (poisoning and shoot- ing) are prohibited because of the endan- gered classification of this species; man has little or no influence upon three (predation, drought, and disease); and only one of the factors (habitat change) is readily amenable to managerial control. Efforts to transplant Utah prairie dogs onto sites of former colonies have had limited suc- cess. Elmore and Workman (1976:21) stated: "In nearly all historic dogtowns, with few ex- ceptions, sagebrush height and density is the restricting factor for any further reintroduc- tion of the animals." This paper presents the results of a study designed to determine if the success of transplanting Utah prairie dogs onto the site of a historic dogtown could be increased by manipulating the vegetation pri- or to the reintroduction of the animals. Study Area and Methods The study was conducted from 1978-1981 at the site of a former colony of Utah prairie dogs located on Jones Bench in the extreme northwest corner of Capitol Reef National Park in south central Utah. Jones Bench lies within a 25-31 cm precipitation belt, and the elevation is 2200 m. Vegetation on the site was dominated by big sagebrush. Blue grama (Bouteloua gracilis) was second most impor- tant in terms of canopy cover. Other plant species found in abundance on Jones Bench were: goosefoot {Chenopodium leptophyl- him), tumbling orach {Atriplex rosea), scarlet globemallow {Sphaeralcea coccinea), bot- tlebmsh squirreltail {Sitanion hystrix), four- wing saltbush {Atriplex canescens), and Yel- low brush {Chrysothamnus viscidiflorus). Five 5-ha plots were established on Jones Bench. Each plot represented a transplant site. Vegetation measurements were taken on the five sites prior to treatment in 1978, and after treatment in 1979 and 1980. The same measurements were taken on 10 active colo- nies of Utah prairie dogs located on the Awapa Plateau, approximately 35 km south- west of Jones Bench in 1978. These measure- ments were taken to determine differences in vegetal characteristics between occupied and unoccupied colonies. The method of vegeta- tional analysis used was that described by Poulton and Tisdale (1961), modified only to the extent of using metric rather than U.S. standard measurements. Four manipulative treatments were planned. They were rotobeating, railing, her- bicide (2,4-D), and fire. The rotobeating was December 1982 Player, Urness: Utah Prairie Dog 519 Table 1. Percent cover and height of plant life forms for an active Utah prairie dog colony on the Awapa Plateau in 1978, and the Jones Bench transplant sites in 1978 (pretreatment), and 1979 and 1980 (posttreatment). Shnib Forb Gi -ass Percent Percent Percent Percent bare Percent Percent cover Height" cover Height cover Height ground litter rock 1978 Awapa Plateau 14.9 24.7 2.9 13.0 3.1 9.4 56.1 14.2 8.8 Rotobeaten 19.2a ° 48.3a 0.5 5.8 6.6 7.3 50.6 14.3 8.7 1978 Railed 18.8a 57.5a 4.1 18.2 1.1 10.5 57.5 16.0 2.5 Herbicide 17.7a 46.1a 6.6 10.4 2.9 14.7 57.6 11.3 .3.9 Control 25.3a 49.1a 1.3 8.7 1.9 7.3 53.8 13.1 4.6 Rotobeaten 1.8a 14.3a 0.5 8.7 12.5 12.0 40.0 37.3 8.0 1979 Railed 2.2a 20.4a 0.2 4.6 1.0 13.3 71.3 22.0 3.4 Herbicide 4.2a 40.8ab 0.1 6.5 4.0 14.6 60.0 27.9 3.7 Control 21.,5l3 45.7b 0.5 29.0 2.2 15.4 61.8 10.5 3.6 Rotobeaten 6.9a .30.9a 0.1 10.3 4.3 25.4 22.6 50.3 15.7 1980 Railed 13.5ab 41.0a 0.3 20.8 0.7 23.1 43.7 33.6 8.2 Herbicide 6.6a 35.8a 0.1 50.3 1.4 38.2 35.6 47.8 8.4 Control 22.9b 59.4a 0.0 13.0 0.4 16.5 26.8 38.3 11.5 'For shrubs only, within the same year and column, means followed by the same letters do not differ significantly at the 0.05 level. ' ° Mean maximum height in centimeters. accomplished by setting the blades at 10 cm above ground level in order to reduce all vegetation to that height. Railing was accom- plished by bolting four medium gauge rail- road rails together. This resulted in a 3.75 m long set weighing 71.4 kg per m, which is comparable to one heavy gauge rail. The site was dragged twice in opposing directions. Both the railing and rotobeating treatments were carried out in late August 1978. At- tempts to achieve the fire treatment failed because there was insufficient ground cover to carry fire between shrubs. As a result, the fire treatment was dropped from the research plan. The herbicide (2,4-D) was applied by a ground sprayer at the rate of 2.22 kg active ingredient per ha with water (123 1/ha) as a carrier. Production of herbaceous species was esti- mated on all transplant sites in August 1979, July 1980, and August 1981. A double- sampling scheme, utilizing a 0.89 m^ circular plot randomly placed 60 times on each site, was used. Green weight of herbage was ocu- larly estimated by 3 observers with the aver- age of the three recorded per plant. Of the 60 plots, every fourth one was clipped and the actual weights obtained for estimate cor- rection via regression analysis. These samples were then air dried to determine percent moisture. Measurements of the visual obstructions to prairie dogs were taken on each transplant site in 1979, 1980, and 1981, and on the site of an active colony on the Awapa Plateau in 1979. A modified version of the technique described by Jones (1968) was used. The method consists of a cover board measuring 65 X 65 cm, with 50 black and 50 white squares each 6.5 X 6.5 cm, arranged in a checkerboard fashion. Thirty readings were taken on each site by randomly placing the board at each site location in one of eight randomly chosen, compass directions. Obser- vations were taken from a height of 30 cm, 20 m from the board. Each site had a max- imal count of 3000 squares visible to the in- vestigator. The ratio of actual number of squares counted to the total possible gave a relative percent visibility for each site. Other characteristics were measured to as- sure homogeneity of the transplant sites. Measurements of soil depth to an impeding layer up to 1 m were taken on each treat- ment site. In addition to this, soil texture and color were determined from a soil sample taken from the surface horizon of each site. The degree of slope was estimated to the nearest five degrees for each site using a hand-held clinometer. The aspect to the nearest 1/8 compass interval was also re- corded for each site. Differences in these characteristics were relatively small. Prior to the actual transplanting, approx- imately 200 artificial burrows, arranged in a matrix, were dug on all sites with a power 520 Great Basin Naturalist Vol. 42, No. 4 Table 2. Visual obstruction measurements (30 observations per location) taken on the transplant sites on Jones Bench in 1979, 1980, 1981, and the site of an active colony of Utah prairie dogs on the Awapa Plateau in 1979. Location Mean percent visibility Number of zero readings Range in percent visibility Control Herbicide Railing Awapa Plateau 1979 2.9a ° 7.3a 17.9a 1980 7.1a 14.3a 11.3a 1981 7.8a 10.7a 17..3a 1979 21 16 10 1980 16 11 7 1981 18 13 11 1979 0-31 0-44 0-62 1980 0-41 0-52 0-62 1981 0-34 0-36 0-58 45.5b 0-83 Rotobeatinj 50.8b 40.6b 44.9b 0 1 0 17-90 0-68 14-82 "For percent visibility within the same year, means followed by the same letters do not differ significantly at the 0.05 level. auger. The holes were dug at an angle and were approximately 9 cm in diameter and 60 to 90 cm deep. Torres (1973) reported that only when artificial burrows were dug at an- gles of 10 to 40 degrees was he successful in reestablishing populations of black-tailed prairie dogs in Colorado. Burrows were dug to provide the animals with temporary pro- tection from predators and adequate thermoregulation.^ A total of 200 prairie dogs (50 per site) was transplanted between 16 June and 4 July 1979. The animals were trapped from five colonies located near Loa, Utah. The capture site was the same elevation as the release site. Twenty immature females, 13 immature males, 6 mature males, and 11 mature fe- males were released on each transplant site. One mature male, one mature female, and two immature animals were placed in three separate cages on each site. This was done to determine if temporarily holding them on the site would more likely assure their per- manent location there in contrast to just re- leasing them on each treatment site (Salmon and Marsh 1981). The cages were construct- ed of 1 X 2 inch hardware cloth and mea- sured 46 cm high, 77 cm wide, and 122 cm long. Centrally located in the screened bot- tom of each cage was a 30 X 30 cm hole that was placed over an artificial burrow. Caged animals had free access to water and were fed whole oats and fresh alfalfa daily. All other animals were individually released into artificial burrows located on high relief areas of their respective transplant sites. All sites were monitored daily in 1979 for animal activity during 23 consecutive days following the release of the first animals. Monitoring took place from elevated loca- tions around the perimeter of the transplant sites. The observer approached close enough to alert the animals (which caused them to stand erect, thus making them more visible), but not so close that they became alarmed and went below ground. Monitoring con- sisted of taking counts during a 10-minute time period on each transplant site during the morning. Biweekly monitoring began after 23 days of daily monitoring. This involved taking the same counts but on two consecutive days every other week throughout the summer and early fall of 1979. In 1980 counts were taken 12-13 June and 21-22 July, and in 1981 counts were taken on 1 July (p.m.), 2 July (a.m.), and 5 August (a.m. and p.m.). The highest count obtained for each transplant site during each observation period was used in a randomized block design for evaluating the relative success of the individual trans- plant sites. The blocks were timed so the var- iance due to time was eliminated from the evaluation. Through this method it was pos- sible to determine if significant differences occurred between the transplant sites. When significant differences did occur, multiple 'David F. Balph, Professor, Department of Wildlife Science, Utah State University, personal interview, 14 March 1978. December 1982 Player, Urness: Utah Prairie Dog 521 Table 3. Grass and forb production, and percent moisture at the transplant sites on Jones Bench for 1979, 1980, and 1981. Transplant site Percent moisture Herb; ige production" ° 1979 1980 1981 1979 1980 1981 Railed Herbicide Rotobeaten Control 50a ° 46a 45a 31a 50a 55a 37a 22a 25ab 49bc 30ab 26a 151c 58a 157d 69b 295b 669c 432b 68a 203b 574b 317b 29a "Within a column, means followed by the same letter do not differ significantly at tl-.e 0.05 level for herbage production and 0.10 for percent moisture. '"Dry weight in kg/ha. comparisons were made using the LSD test of Fisher (Ott 1977). Results Percent cover and height of plant life foniis were considered more important than any particular botanical composition because prairie dogs are opportunists that eat any available forage that has nutritional value (Koford 1958, Crocker-Bedford 1976). There- fore, the plant species were grouped accord- ing to their life form. Pretreatment shrub height on Jones Bench is the only vegetal measurement taken that showed a significant difference (at the 0.01 level) from that of the shrub height of active colonies. This strength- ened the assumption that the pretreatment vegetational height on Jones Bench was too tall for successful transplanting of prairie dogs. The different treatments had highly varied effects upon the vegetation (Table 1). Per- cent cover of shrubs, primarily big sagebrush, was the only characteristic that was greatly reduced by all treatments. Shrub cover on manipulated sites differed significantly from the control, with the exception of the railed area the second year after treatment (1980). Shrub height was reduced by railing and rotobeating the first year following treatment (1979), but was not significantly reduced on the herbicide treatment because skeletons of dead plants remained intact. There were no significant differences in shrub height in the second year posttreatment because of rapid recovery of shrubs on the rotobeaten and railed sites. Shrub height on the herbicide area continued to decline slowly as dead plants disintegrated. Table 2 shows the visual obstruction mea- surements. With all comparisons (in all years) 'Some animals moved to locations near but off the designated transplant sites. only those taken on the rotobeaten site do not differ significantly when compared to measurements taken on the site of an active prairie dog colony on the Awapa Plateau. Although the percent moisture of the herb- age varied greatly between transplant sites because of wide variability among the mea- surements taken within each site (Table 3), significant differences (at the 0.10 level) were found only in 1981. There were significant differences, however (at the 0.05 level), in the total herbage production between the transplant sites. In 1979 all sites differed sig- nificantly from one another. In 1980 only the railing and rotobeating sites did not differ significantly, and in 1981 all sites differed sig- nificantly from the control. Table 4 lists results of the animal counts taken during 1979, 1980, and 1981. In 1979 the rotobeating site had significantly higher numbers of animals than the other sites; there were no significant differences between the railed and other sites^ or between the herbi- cide and control sites. In 1980 and 1981 no animals were observed on either the control or herbicide sites. In 1980 and 1981 the roto- beaten and railed sites did not differ signifi- cantly, but they did differ significantly from all other sites. To a certain extent, prairie dogs were more easily seen on the more open treatments, and this may have affected the counts somewhat. However, the less visible sites were carefully checked for signs of fresh diggings; when such signs were found, these areas were more closely observed. Discussion Of the animals placed in cages in an at- tempt to get them to locate at the release site, all the adults had dug out of their cages within five days; one adult male dug out in 522 Great Basin Naturalist Vol. 42, No. 4 Table 4. Mean numbers of animals coimted on Jones Bench during 1979, 1980, and 1981. Count sites Year Rotobeaten Railed Other' Herbicide Control 1979 (n = 9)' 1980 (n = 2) 1981 (n = 2) 16.0c 13.5b(5)^ 15.0b(9) 8.1b^ 8.5b(3.5) 15.5b(12) 3.,3ab 1.0a 2.0a(l) 1.7a 0.0a 0.0a 0.3a 0.0a 0.0a 'Some animals moved to sites on Jones Bench other than designated transplant sites. 'Within the same year, means followed by the same letter do not differ significantly at the 0.05 level. 'Number of counts taken during the year. 'Mean number of young counted are in parentheses. less than three hours. On the rotobeaten and railed sites these or other animals occupied some of the cages and their underlying tun- nels throughout the first summer. Immature animals were much slower in digging out; some did not dig at all for nine days, so they were released. It is doubtful that immature prairie dogs could survive if they were re- leased in areas without burrows or adult ani- mals to dig burrows. The longevity of vegetational treatments is related to the amount of brush removed by the treatment (Nielsen and Hinckley 1975). The rotobeaten and railed sites will require retreatment every five to ten years. The her- bicide treatment would likely not require such a short retreatment period. In 1981 there was evidence that animals may be mov- ing onto the herbicide site. This is likely be- cause, while other treatments are returning to their pretreatment state, the herbicide is becoming more favorable as habitat. The skeletal remains of the herbicide-killed shrubs are deteriorating; thus visibility for prairie dogs is increasing. It may be possible to greatly reduce the need for retreatment by combining vegeta- tion treatments. If rotobeating were to be fol- lowed in the next year or two by spraying with 2,4-D, then a higher percent kill of shrubs could be attained, as well as an effec- tive reduction of visual obstructions. Treat- ment could, of course, follow the reverse se- quence for the same effect. The controlled use of fire may be the best technique to achieve the desired results where fuel loading is sufficient to allow burn- ing. Fire, if carried out properly, could re- move a high percentage of nonsprouting shrubs and increase visibility immediately at low cost. With such results it is likely that re- treatment would not be necessary for per- haps 20 years or more. Fire would also re- lease many grasses and forbs for increased growth, thus making the site even more fa- vorable for prairie dog reestablishment through increased food resources. The negative response of transplanted ani- mals to the control site was a strong in- dication that some type of vegetal treatment is necessary when transplanting animals onto sites of former colonies presently overgrown with shrubs. The chances of a successful transplant could be increased by first reduc- ing shrub height and cover. Our study should aid in reestablishing scattered colonies of Utah prairie dogs throughout their former range to help assure the continued existence of this unique animal. One objective of this effort is to restore sufficient healthy popu- lations on public lands to allow for delisting of this animal as an endangered species and thus reduce conflicts on private lands by per- mitting local control on agricultural problem areas. Literature Cited Allan, P. F., and B. Osborn. 1949. Vegetation of an abandoned prairiedog town in tall grass prairie. Ecology 30:322-332. Blaisdell, J. p. 1953. Ecological effects of planned burning of sagebrush: grass range on the upper Snake River Plains. USD A Tech. Bull. 1075. 39 pp. Bureau of Sport Fisheries and Wildlife. 1968. Rare and endangered fish and wildlife of the United States. U.S. Govt. Print. Off., Washington, D.C. 153 pp. 1972. Rare and endangered fish and wildlife of the United States. U.S. Govt. Print. Off., Wash- ington, D.C. 272 pp. 1973. Rare and endangered fish and wildlife of the United States. U.S. Govt. Print. Off., Wash- ington, D.C. 289 pp. Collier, G. D. 1974. The Utah prairie dog: abundance, distribution, and habitat requirements. Unpub- December 1982 Player, Urness: Utah Prairie Dog 523 lished dissertation. Utah State Univ., Logan. 96 pp. Collier, G. D., and J. J. Spillett. 1972. Status of the Utah prairie dog (Cynornys parvidens). Utah Acad. Sci., Arts, and Lett. 49:27-39. 1973. The Utah prairie dog— decUne of a legend. Utah Science 34:83-87. Crocker-Bedford, D. C. 1976. Food interactions be- tween Utah prairie dogs and cattle. Unpublished thesis, Utah State Univ., Logan. 118 pp. Crocker-Bedford, D. C, and J. J. Spillett. 1977. Home ranges of Utah prairie dogs. J. Mammal. 58:63. CoTTAM, W. P., AND F. EvANS. 1945. A Comparative study of the vegetation of grazed and ungrazed canyons of the Wasatch Range, Utah. Ecology 26:171-181. Ellison, L. 1960. Influences of grazing on plant succes- sion of rangelands. Bot. Rev. 26:1-78. Elmore, S., and G. Workman. 1976. A baseline study of the past and present status of the Utah prairie dog {Cynomys parvidens) in Bryce Canyon Na- tional Park. Utah State Univ., Dept. Wildl. Sci., Logan. 40 pp. Fitzgerald, J. P., and R. R. Lechleitner. 1974. Obser- vations on the biology of Gunnison's prairie dog in central Colorado. Am. Midi. Nat. 92:142-163. Jones, R. E. 1968. A board used to measure cover used by prairie grouse. J. Wildl. Manage. 32:28-31. Koford, C. B. 1958. Prairie dogs, white faces and blue grama. Wildl. Monog. 3:1-78. Nielsen, D. B., and S. D. Hinckley. 1975. Economic and environmental impacts of sagebrush control on Utah's rangelands. A review and analysis. Utah Agric. Expt. Sta. Res. Rept. No. 25. 27 pp. OsBORN, B. 1942. Prairie dogs in shinnery (oak scrub) savannah. Ecology 23:322-332. Ott, L. 1977. An introduction to statistical methods and data analysis. Duxbury Press, North Scituate, Massachusetts. 730 pp. PicKFORD, G. D. 1932. The influences of continued heavy grazing and promiscuous burning on spring-fall ranges in Utah. Ecology 13:159-171. PouLTON, C. E., AND E. W. TiSDALE. 1961. A quan- titative method for the description and classifica- tion of range vegetation. J. Range Manage. 14:13-21. Salmon, T. P., and R. E. Marsh. 1981. Artificial estab- lishment of a ground squirrel colony. J. Wildl. Manage. 45(4): 1016-1018. Schaffner, J. H. 1929. Extension of the natural range of two mammals in Clay County, Kansas. Trans. Kansas Acad. Sci. 31:61-62. Smith, A. D. 1949. Effects of mule deer and livestock upon foothill range in northern Utah. J. Wildl. Manage. 13:421-423. Snell, G. p., and B. D. Hlavachick. 1980. Control of prairie dogs— the easy way. Rangelands 2(6):239-240. Torres, J. R. 1973. Status of the black-tailed prairie dog and the black-footed ferret in North Dakota. Pages 51-59 in Proceedings of the Black-Footed Ferret and Prairie Dog Workshop. South Dakota State Univ., Brookings. 208 pp. TuELLER, p. T., and W. H. Blackburn. 1974. Condition and trend of the big sagebrush /needle-and-thread grass habitat type in Nevada. J. Range Manage. 27:36-40. EFFECTS OF DEFOLIATION ON REPRODUCTION OF A TOXIC RANGE PLANT, ZIGADENUS PANICULATUS V. J. Tepedino' Abstract.— The effect of complete defoliation, prior to flower stalk appearance, on the reproductive success of foothill deathcamas, a toxic range plant, was studied in northern Utah. Defoliated plants did not replace their leaves. Defoliation had no effect on total number of flower stalks produced but did significantly slow the rate of stalk emer- gence and reduce the number of plants that produced open flowers. The number of leaves produced bv control plants was also positively associated with the probability of producing a flowering stalk. Few plants in either defo- liated or control treatments set seed, probably because of inactivity of pollinators during a cold and wet spring. It is suggested that species, such as deathcamas, which either produce leaves early in spring or are liliaceous geophytes, may be especially vidnerable to herbivory. Among the characteristics thought to ren- der plant species "apparent," or relatively easy for herbivores to find, are the peiennial habit and large size, both of individuals and of populations (Feeney 1976, Rhoades and Cates 1976, Rhoades 1979). Apparent species are also hypothesized to reduce their suscep- tibility to herbivory by diverting relatively large amounts of energy from vegetative and reproductive functions to the production of antiherbivore compounds. Foothill deathcamas (Zigadeniis pan- iculatus [Nutt] S. Wats; Liliaceae) is a bulb- forming range plant of the western U.S. (James et al. 1980) that possesses some of these characteristics of apparency: it is pe- rennial and is commonly found in large num- bers throughout its range, although individual plants are small. Apparency is further in- creased because deathcamas is among the first species to produce leaves in the spring (James et al. 1980, pers. obs.): it is therefore extremely attractive to mammalian herbi- vores that have subsisted on low-quality for- age through the winter. Despite their avail- ability, few plants {^ 10.0 percent) display any evidence of herbivory (Tepedino, un- publ. ms.), evidently because of the numerous steroid alkaloids present in the leaves and other plant parts (Willaman and Li 1970) that are toxic to mammals (Marsh and Claw- son 1922). In addition to the apparently large com- mitment to the defense of leaf tissue by the production of alkaloids, there are other rea- sons for suspecting deathcamas to be particu- larly vulnerable to herbivory when it occurs. Although most perennials typically replace their leaves soon after defoliation (Jameson 1963, Kulman 1971, Rockwood 1974), evi- dence suggests that geophytic species of the Liliales may be incapable of doing so (Heath and Holdsworth 1948); under normal condi- tions, once the presumptive flower stalk bud is formed in early spring no further leaf ini- tials are cut. Also, a minimum number of leaves has been shown to be necessary for flower stalk production for several liliaceous species (Heath and Holdsworth 1948 and ref- erences therein). Thus, defoliation of death- camas may significantly impair reproductive success by lowering the number of flowers or seeds produced. The relationship between the number of leaves and flowers produced and the effect of artificial defoliation on re- productive success are reported here. Methods The study site (alt. 1400m) was at the top of a west-facing embankment, 8 km south of Avon, Utah (Cache Co.), along County Road 165. Here numerous deathcamas plants grew among sagebrush (Arteinisia sp.) and associ- ated forbs. 'USDA-ARS-WR, Bee Biology and Systemalics Laboratory, Utah State University, UMC 53, Logan, Utah 84322. 524 December 1982 80 Tepedino: Deathcamas Reproduction STALKS b FLOWERS 525 (O 70 60 < 50 -J a. S « UJ 30 00 I 20 10 5/1 6 11 16 21 26 31 6/510 5/1 6 11 16 21 26 31 6/5 10 DATE Fig. 1. Number of plants producing a, flowering stalks; b, open flowers for defoliation (dashed line); and control (solid line) treatments. Foothill deathcamas produces 3-9 basal leaves from a tunicate bulb in early spring, followed by a single paniculate flowering stalk 4-6 weeks later. Plants with 5 or more leaves were completely defoliated on 23 April 1981 after the basal leaves were fully extended, but before appearance of the flowering stalk. Leaves of 100 plants were counted and removed at soil level with a ra- zor blade. At the same time, a plant nearby (between 1-2 m) each of those treated was selected as a control. Leaves of controls were also counted. All plants were marked with plastic labels. Plants were subsequently examined at ap- proximately weekly intervals for devel- opmental stage of the flowering stalk. Ab- sence or presence of the stalk, presence of open or spent flowers, and fruit maturation (judged by expansion of the perianth) were recorded. Precipitation and temperature records were obtained from a Utah State University weather station located 19 km north of the site at similar elevation. FIesults Unlike some other perennial species, Z. paniculatus did not replace lost leaf tissue; once defoliated, experimental plants re- mained in that condition for the rest of the growing season. Flowering stalks of plants from both treat- ments had begun to emerge by the first ex- amination date (1 May). However, the rate of emergence was slower for defoliated plants than for controls (Fig. la); a chi-square test of equal distribution of plants with flowering stalks in each treatment was significant for the first three sampling dates (1 May, X^ = 3.86, P = 0.05; 7 May, X2 = 7.58, P<0.01; 12 May, X2 = 3.97, P = 0.05). Although flowering stalks of defoliated plants emerged more slowly, by the end of the flowering sea- son there was no significant difference be- tween treatments in the total number of stalks produced (X^ = 0.35, P<0.50). The major difference between defoliated and control plants was in the number of flowering stalks that produced open flowers 526 Great Basin Naturalist Vol. 42, No. 4 (Fig. lb). Only 21 of the 66 stalks produced by defoliated plants developed open flowers as compared to 59 of 71 control stalks. (Two additional control stalks were decapitated by herbivores before they flowered). Flower buds on most stalks of defoliated plants with- ered soon after the stalks emerged, and stalks subsequently turned brown. Few plants of either treatment set seeds. Only one defoliated plant, and five control plants produced seeds. Lack of seed set by control plants was probably due to inactivity of pollinators caused by cold and rainy weather. Precipitation fell on 15 of 31 days in May and the maximum temperature was below 21 C (70 F) on 18 days. Only 7 days were both rainfree with maximum temper- ature at or above 21 C, and all these came at the beginning or end of the month (days 1, 2, 25, 28-31). The number of leaves produced by a plant was associated with stalk production for both treatments. Table 1 shows the distribution of plants by leaf number categories; the overall distribution did not differ significantly be- tween treatments (X^ = 1.38, d.f. = 3; in this and the following analysis leaf number categories have been combined when ex- pected values were <5.0, Maxwell [1961]). For each treatment the proportion of plants producing stalks increased with leaf number (Table 1). A comparison of the number of plants in each leaf category that produced stalks with those that did not was significant for both treatments (defoliated, X^ = 7.2, d.f. = 2, P<0.05; control, X2 = 12.2, d.f. = 1, P<0.001): plants with more leaves had a greater probability of producing a stalk even when leaves had been removed. A com- parison between treatments of the distribu- tion of plants with stalks by leaf number showed no significant difference (X^ = 0.06, df. = 2, P<0.50). Plants with more leaves also exhibited a greater tendency to produce open flowers (Table 1). However, when plants producing flowers were compared by leaf number with plants producing only stalks, differences be- tween treatments were apparent. For control plants the distributions did not differ signifi- cantly (X2 = 0.96, d.f. = 1); if a plant sent up a stalk, there was a high probability (80.8 percent) that flowers would open, irrespec- tive of leaf number. For defoliated plants there was a greater probability for plants that had produced more leaves to produce some open flowers on the stalk than for those with fewer leaves (X2 = 3.36, d.f. = 1, P = 0.07). Discussion Defoliation had irreparable effects on the reproductive potential of Z. paniculatus. Al- though removal of leaves just before stalk emergence did not affect the likelihood of producing a stalk, it did significantly delay the emergence of stalks and reduce the prob- ability that any flower buds would reach an- thesis. These results are in general agreement with other studies that have shown that simu- lated herbivory can significantly reduce num- bers of flowers (Blaisdell and Pechanec 1949, Callan 1949, Simmonds 1951, Mueggler 1967, Enyi 1975) or seeds (Sackston 1959, Rockwood 1974, Enyi 1975), and also delay flowering (Collins and Aitken 1970) (see Jameson 1963, Kulman 1971 for reviews). The effects of defoliating deathcamas, however, may be more profound than the simple elimination of a single year's repro- duction. The results suggest that stored car- bohydrates from the previous year are de- pleted in the production of leaves. Leaves, in turn, hasten emergence of the stalk, and are required for maturation of flowers, seeds, and the synthesis of storage material for the sub- sequent year's vegetative growth. But, unlike many other perennials, Z. paniculatus is ap- parently unable to produce a second crop of leaves after defoliation. If leaves are cropped Table 1. Distribution of defoliated and control plants by number of leaves/plant, and the percentage of total plants in each category that produced flowering stalks and open flowers. Number of leaves 5 6 7 8 9 Defoliated Total no. With stalk (%) With flowers (%) Control Total no. With stalk (%) With flowers (%) 0 0.0 0.0 2 0.0 0.0 13 38.5 15.4 8 25.0 25.0 53 64.2 13.2 60 70.0 53.3 29 75.9 34.5 24 95.8 83.3 5 100.0 40.0 6 100.0 83.3 December 1982 Tepedino: Deathcamas Reproduction 527 before stalk emergence, plants may have had insufficient photosynthetic surface to pro- duce storage material for the following year. Thus, it is possible that a single defoliation episode is sufficient to cause either death of the plant or to eliminate reproduction for more than one year. If this proves to be the case, the production of antiherbivore com- poimds becomes important. The positive relationship between the nmnber of leaves produced and the likeli- hood of sending up a flowering stalk has also been reported for other species in the Liliales (Heath and Holdsworth 1948). These results suggest that the number of leaves produced increases with age of the plant, and that plants do not begin flowering until the sec- ond or third year. This assertion needs more careful examination; however, numerous plants with 3-5 leaves in a population at higher altitude (1850 m) have been observed both to bloom and produce seeds (Tepedino, unpubl. ms.). Schemske et al. (1978) showed that seed production in self-incompatible spring- flowering herbs can vary considerably from year-to-year, and suggested that this variation was due to the effect of unpredictable spring weather on pollinators. A similar explanation seems appropriate for Z. paniculatus, which also requires insects for pollination (Tepedino 1981). Few plants in the control treatment set seed, and this was associated with an ab- sence of pollinators during an extended peri- od of cold and wet weather. Conversely, in the previous year, when weather was more conducive to insect activity, most plants set seed at a nearby site (pers. obs.). Two characteristics of the growing season and life form of Z. paniculatus suggest pos- sible modifications of the concept of plant apparency. First, as noted above, plant spe- cies that leaf out early in spring are extreme- ly apparent to mammalian herbivores, and we should expect the leaves of such plants to be well defended. This seasonality com- ponent of apparency seems to have been pre- viously ignored. A cursory perusal of James et al. (1980) suggests that 75-80 percent of the plants most poisonous to livestock in the western U.S. begin growth in early spring, so the idea would seem to deserve further attention. Second, several workers have noted that, among the monocots, only the Liliales are well represented in the number of species that produce alkaloids (Hegnauer 1966, Levin and York 1978). Tomlinson (1980) has pointed out that many of these species are geophytes. Perhaps the leaves of many of these species are irreplaceable, as in death- camas and other liliaceous species (Heath and Holdsworth 1948), and must therefore be heavily defended against herbivory. Acknowledgments I thank K. Ruggeri, T. Peery, and M. Schultz for help with the field work and data tabulation; T. Waldron for inking the figure; and D. R. Frohlich and Drs. E. Bach Allen and I. Palmblad, Utah State University, for commenting on the manuscript. Literature Cited Blaisdell, J. p., AND J. P. Pechanec. 1949. Effects of herbage removal at various dates on vigor of bluebunch wheatgrass and arrowleaf balsamroot. Ecology 30:298-305. Callan, E. M. 1949. Effect of defoliation on reproduc- tion of Cordia inacrostacbifa. Bull. Entomol. Res. 39:213-15. Collins, W. J., and Y. Aitken. 1970. The effect of leaf removal on flowering time in subterranean clo- ver. Aust. J. Agric. Res. 21:893-903. Enyi, B. a. C. 1975. Effects of defoliation on growth and yield in groundnut {Arachis hypogea), cowpeas {Vigna unquiculata), soybean {Glycine max), and green gram (Vigna aureus). Ann. Appl. Biol. 79:55-66. Feeny, p. 1976. Plant apparency and chemical defense. Rec. Adv. Phytochemistry 10:1-40. Heath, O. V. S., and M. Holdsworth. 1948. Morpho- genic factors as exemplified by the onion plant. Sym. Soc. Exp. Biol. 2:326-350. Hegnauer, R. 1966. Comparative phytochemistry of al- kaloids. Pages 211-230 in T. Swain, ed.. Com- parative phytochemistrv. Academic Press, New York. James, L. P., R. F. Keeler, A. E. Johnson, M. C. Williams, E. H. Cronin, and J. D. Olsen. 1980. Plants poisonous to livestock in the western states. USDA Agric. Infor. Bull. 415. 90 pp. Jameson, D. A. 1963. Responses of individual plants to harvesting. Bot. Rev. 29:532-594. Kulman, H. M. 1971. Effects of insect defoliation on growth and mortality of trees. Annu. Rev. Ento- mol. 16:289-324. Levin, D. A., and B. M. York, Jr. 1978. The toxicity of plant alkaloids: an ecogeographic perspective. Biocheni. Svst. Ecol. 6:61-76. 528 Great Basin Naturalist Vol. 42, No. 4 Marsh, C. D., and A. B. Clawson. 1922. The death- camas species, Zijgadeniis paniculatiis and Z. ele- gans, as poisonous plants. USDA Bull. No. 1012. 25 pp. Maxwell, A. E. 1961. Analysing qualitative data. Methuen, London. MuEGGLER, W. F. 1967. Response of mountain grassland vegetation to clipping in southwestern Montana. Ecology 48:942-949. Rhoades, D. F. 1979. Evolution of plant chemical de- fense against herbivores. Pages 3-54 in G. A. Ro- senthal and D. H. Janzen, eds.. Herbivores: their interaction with secondary plant metabolites. Ac- ademic Press, New York. FIhoades, D. F., and R. G. Gates. 1976. Toward a gen- eral theory of plant antiherbivore chemistry. Rec. Adv. Phytochemistry 10:168-213. RocKwooD, L. L. 1974. The effect of defoliation on seed production of six Costa Rican tree species. Ecolo- gy 54:1363-1.369. Sackston, W. E. 1959. Effects of defoliation on sun- flowers. Can. J. Plant Sci. 39:108-118. Schemske, D. W., M. F. Willson, M. N. Melampy, L. J. Miller, L. Verner, K. M. Schemske, and L. B. Best. 1978. Flowering ecology of some spring woodland herbs. Ecology 59:351-66. SiMMONDS, F. J. 1951. Further effects of the defoliation of Cordia macrostachi/a (Jacq.) R. and S. Cana- dian Entomol. 83:24-27. Tepedino, V. J. 1981. Notes on the reproductive biology of Zigadenus paniculatiis, a toxic range plant. Great Basin Nat. 41:427-430. ToMLiNSON, P. B. 1980. Monocotyledonous habit in rela- tion to morphology and anatomy. Sym. Linn. Soc. London 8:7-19. WiLLAMAN, J. J., AND H. L. Ll 1970. Alkaloid-bearing plants and their contained alkaloids. Lloydia (supplement) 33 (3 A). 286 pp. DISTRIBUTION AND RELATIVE ABUNDANCE OF FISH IN RUTH RESERVOIR, CALIFORNIA, IN RELATION TO ENVIRONMENTAL VARIABLES' Steven Viifg- and Thomas J. Hassler' Abstract.— The fish population of Ruth Reservoir, Cahfornia, was sampled every two weeks with variable mesh gill nets from May 1974 through May 1975. Fish were captured in the following order of numerical abundance: Humboldt sucker (Catostomus hurnboldtianus), golden shiner {Notemigonus crysoleucus), brown bullhead [Ictahtrus nebulosus), white catfish (/. catus), rainbow trout {Sahno gairdneri), and largemouth bass {Micwpterus sabnoides). The three most abundant species made up about 95 percent of total numbers and weight. All species exhibited a similar cyclic temporal availability pattern: catch rates increased to a maximum during summer and fall and de- creased during winter and spring. Environmental variables with the most pronounced relationships to fish catches were temperature (direct) and turbidity (inverse). Information on Ruth Reservoir fish ecology collected prior to this study was limited; data consisted of stocking records, yearly creel survey data on opening weekends of the fish- ing season, five gill net sets during November 1968, and the results of a reward tagging pro- gram for salmonids during May 1972 (Ruth Reservoir file, California Department of Fish and Game, Eureka). Management measures have consisted primarily of stocking hatch- ery-reared salmonids. Unauthorized in- troductions of exotic species into the reser- voir have also been made. Alterations to the dam have been proposed that would affect the physical and chemical characteristics of the lake and thus the aquat- ic organisms, specifically the fish populations. The present dam may be modified or re- placed by a larger structure to meet future water needs (U.S. Army Corps of Engineers 1973). Air-induced circulation and a multi- level discharge structure have been proposed to reduce downstream turbidity (Winzler and Kelly 1975). The objectives of this study were to deter- mine the relative abundance and distribution of fish in Ruth Reservoir and determine their relation to environmental variables. Study Area Ruth Reservoir is impounded behind R. W. Matthews Dam, near the headwaters of the Mad River in Trinity County, California (Fig. 1). This water supply reservoir, about 127 km by river from the Pacific Ocean, provides municipal and industrial water for the Hum- boldt Bay Area. The dam was completed in 1961 and is operated by Humboldt Bay Mu- nicipal Water District (HBMWD). The reser- voir has a maximum surface area of 445.2 ha, a maximum storage capacity of 63.9 million cubic meters, a mean depth of 14.4 m at maximum pool, and a minimum discharge of 142 liters per second. Annual water level fluctuations have ranged from 9.8 to 15.5 m, with a mean fluc- tuation of 12.6 m (HBMWD, unpublished data). The water level is usually lowest in November and highest in January. The high- est recorded water level, 5.8 m above spillway elevation, was on 22 December 1964, and the lowest, 13.7 m below spillway elevation, on 29 November 1967. Water level fluctuated 14.4 m during the study. The reservoir is 11.3 km long at full pool and has a mean width of 0.6 km (Winzler and 'Cooperators include the California Department of Fish and Game and the United States Fish and Wildlife Service. 'Bioresources Center, Desert Research Institute, P.O. Box 60220, Reno, Nevada 89506. 'California Cooperative Fishery Research Unit, Humboldt State University, Areata, California 95521. 529 530 Great Basin Naturalist Vol. 42, No. 4 ! N California DAM RUTH RESERVOIR SCALE KILOMETERS Fig. 1. Gill net sampling stations in Ruth Reservoir, California MAD RIVER Kelly 1975). Several small tributaries flow into the reservoir, but the major inflow is from the Mad River, which has a watershed of 30,822 ha above the dam (Iwatsubo et al. 1972). The dominant geological feature of the watershed is the Franciscan Formation. Heavy precipitation, steep slopes, and un- stable geology have resulted in high erosion in this area (Yoimg 1971). The primary influx of sediment corresponds to major precipi- tation from November through April. During the rainy season, large amoimts of fine sus- pended sediment are distributed throughout the mixed reservoir; surface turbidity dimin- ishes by late spring as the suspended particles settle and thermal stratification confines sus- pended sediments to the bottom zone, thus developing a turbid density current (Winzler and Kelly 1975). Persistent turbidity occurs in the reservoir and downstream from the reservoir's bottom discharge. Surface water temperatures range from 0 to 26.7 C; the minimum generally occurs in December or January, and the maximum in July or August. Bottom temperatures range from less than 4 to as high as 17 C. The reser- voir has characteristic spring and fall over- turns of a dimictic lake. Dissolved oxygen December 1982 ViGG, Hassler: Ruth Reservoir Fish 531 (DO) ranges from saturation to seasonal an- aerobic bottom deficits (California Depart- ment of Water Resources, Red Bluff, 1969). Methods Sampling was conducted at two-week in- tervals from May 1974 to May 1975; 26 sam- ples—7 each in summer and fall and 6 each in winter and spring— were analyzed by season. The seasons were defined as follows: summer, 1 June to 31 August; fall, 1 September to 1 December; winter, 2 December to 2 March; and spring, 3 March to 31 May. Five gill net sampling stations were established from the dam to reservoir headwaters (Fig. 1). Fish populations were sampled with bot- tom set, variable mesh gill nets 1.83 X 54.86 m comprising six 9.14-m panels of the follow- ing mesh sizes (bar measure): 1.27, 1.91, 2.54, 3.18, 3.81, and 6.35 cm. All mesh sizes were made of nimiber 104 multifilament white ny- lon except the 6.35-cm mesh, which was number 139. The nets were set in the late evening and fished overnight for 12 to 16 hours. Fish catch was adjusted to a standard 12-hour set. The net was anchored in approximately 2 m of water at the inshore end and set per- pendicular to shore. The end of the net placed closest to shore was randomized. Each gill net panel was marked with a painted ver- tical stripe to give two replicates for each set. Fish catch from the right and left halves of each mesh size was recorded separately and randomly assigned to one of two derived replicates. The data could thus be treated as replicate 27.43 m variable mesh nets in each location at each time, enabling the use of a nested analysis of variance design. Limnological data were obtained during each sampling period at each station. Tem- perature, turbidity, conductivity, and DO were measured at limnetic stations corre- sponding to the gill net stations. Water sam- ples were taken with a 2-1 water bottle, 1 m below the surface, at middepth, and 1 m above the bottom. Immediately upon bring- ing the sample to the surface we measured temperature with a mercury bulb thermome- ter, or the thermistor of the DO meter. A bathythermograph was used to measvire depth-temperature profiles. Turbidity, Jack- son Turbidity Units (JTU), was measured with a Hach Model 1860 Turbidimeter. A Beckman Solu Bridge was used to measure electroconductivity, recorded as micro mhos per centimeter (/imho/cm) at 25 C. Dissolved oxygen determinations were made with a Hach Model CA-10 DO kit (June through October) and a Delta Scientific Model 85 DO Meter (November through May). Surface and discharge temperature, reservoir surface ele- vation, inflow, and discharge data were ob- tained from HBMWD records. To detect significant differences in hori- zontal and seasonal fish distribution, and in- teraction between reservoir area and season, we analyzed the catch-per-unit-of-effort data by using a two-way nested analysis of vari- ance design computer program. Fish relative abundance was analyzed by sampling station and season. One-way analysis of variance (Sokal and Rohlf 1969) was used to analyze seasonal differences in mean fish catch at Sta- tion 5. This station was dewatered by sea- sonal low water and was not included in the overall analysis. Fish catch data were transformed: (logio(Y + 1)) where Y = fish catch. Table 1. Gill net catches at five sampling stations in Ruth Reservoir from May 1974 through May 1975. Species Relat ive abundance Relative Mean biomass (w Total eight, g) Number Percent of total Percent of Common name Scientific name Catch /h total Humboldt sucker Catostomtts hiimboldtianus 1.06 1,854 42.5 452.0 838,008 76.3 Golden shiner Notem igon us cn/soleucits 0.82 1,4.32 32.8 53.8 77.042 7.0 Brown bullhead Ictalurtis nebitlosus 0.51 888 20.3 149.9 133,111 12.1 White catfish I. cattis 0.08 141 3.2 262.5 .37,013 3.4 Rainbow trout Sahno gairdneri 0.02 28 0.6 258.0 7,224 0.7 Largemouth bass Microptertis sahnoides 0.01 23 0.5 276.0 6,348 0.6 Total 2.49 4,366 100 - 1,098,746 100 532 Great Basin Naturalist Vol. 42, No. 4 CUBIC REGRESSION EQUATION X*-l-0. Y=-IOI.2IO-h79.743X -6.477X -1-0. 14 1 X {r^= ..792) O CALCULATED VALUES • ACTUAL DATA 300 I WINTER I SPRING 200 I 13 24 9 23 JUNE JULY 1974 Fig. 2. Relationship between time and total fish catch in Ruth Reservoir, June 1974 through May 1975. 6 23 4 19 5 19 2 16 30 14 30 II 25 15 1 15 26 13 26 10 24 AUG. SEPT OCT NOV. DEC. JAN. 1975 FEB. MAR. APRIL MAY Correlation analysis was used to determine relationships between environmental factors and fish catch (by species and station). Prod- uct-moment correlation coefficients (r) be- tween environmental parameters and fish catch were determined and tested for signifi- cance. We included variables with mean- ingful biological relationships that were sig- nificantly correlated with fish catch in multiple linear correlation analyses using the method of least squares (Cooley and Lohnes 1971). Results Species Composition and Relative Abundance Humboldt sucker, Catostomus humbold- tianus, the most abundant of the six fish spe- cies in the net catches, accounted for over 42 percent of the numbers of fish and over 76 percent of the weight (Table 1). Numerically, golden shiner, Noteinigonus crysoleucus, was second in abundance and brown bullhead, Ic- talurus nebulosus, third. Over 1.5 times as many shiners were captured as bullheads; however, the biomass of brown bullheads was over 1.7 times greater than that of golden shiners. White catfish, Ictalurus catiis, made up slightly over 3 percent in terms of num- bers and weight. Largemouth bass, Micro- ptenis salmoides, and rainbow trout, Sahno gairdneri, accounted for less than 1 percent of the catch. Humboldt suckers and rainbow trout were the only fish captured in Ruth Reservoir that were established in the upper Mad River drainage prior to impoundment. The reser- voir has been stocked yearly with rainbow trout from various hatcheries. The coho salmon, Oncorhynchus kisutch, was in- troduced in 1972, and the Japanese ayu, Ple- coglossiis altivelis, in 1964-1965. Neither of these two species was taken in the net catches. Distribution A total of 4366 fish were caught in 1756 gill net hours. Total catch was highest during late summer and early fall, greatly decreased during winter and early spring, and remained low until late spring (Fig. 2). All species showed a cyclic seasonal trend (Fig. 3). The increased catch of golden shiners in the spring sample was most evident, and ac- counted for 52 percent of the total spring catch. Total catch was consistently higher at the upper end of the reservoir (Stations 4 and 5) during all seasons, accounting for 55 to 86 percent of the catch (Table 2). Catches were most evenly distributed during fall, when December 1982 ViGG, Hassler: Ruth Reservoir Fish 533 75 r Humboldt Sucker Golden Shiner Brown Bullhead White Catfish Sumnner Fall Winter Spring Fig. 3. Mean fish catch by species and season in Ruth Reservoir, 1974-1975. about 45 percent of the fish were taken at the lower three stations. Catches of Hum- boldt suckers, golden shiners, and brown bull- heads illustrate this trend of higher abun- dance at upper reservoir stations. Catches of white catfish were more evenly distributed throughout the reservoir; but were slightly higher in the middle and lower than in the upper reservoir areas. Himiboldt suckers were generally more abundant in catches than other species during all seasons. However, during summer and fall catches of brown bullhead were highest. Golden shiner catches were relatively high during summer at Station 5 and during spring at Stations 2 and 4. White catfish were the second or third most abundant species in the catch at the lower end of the reservoir (Sta- tions 1 and 2) during all seasons except win- ter, when only one was captured in the entire lake. Temporal and spatial fish distribution pat- terns were summarized by analysis of vari- ance. Mean catches of Humboldt suckers, golden shiners, brown bullheads, and total species were significantly different (F<0.01) with respect to season and station. The mean catch of white catfish differed significantly by season (?< 0.001) but not by station. A significant interaction for catches of brown bullhead by season and station (P<0.01) in- dicated that seasonal distribution was not consistent on a spatial basis. There was a large difference in seasonal mean catches of bullhead in the upper end of the reservoir, but catches in the lower end were con- sistently low and not greatly different. There was no significant interaction for Humboldt sucker, golden shiner, white catfish, and total catch. Thus, mean seasonal catch for these species was independent of lake area effects and, conversely, differences in area fish catch were significant, regardless of season. There was a significant difference in rep- licates within season and station for all spe- cies (P< 0.005). Seasons were not biologically discrete units of time, i.e., temporal trends of fish catch existed within seasons. At station 5 total species mean catch was significantly higher for the summer-fall than for winter-spring (P< 0.001, Table 3). There was a significant difference in mean catch of 534 Great Basin Naturalist Vol. 42, No. 4 brown bullhead (P< 0.001) and golden shiner (P<0.05) between the two time periods. Catches of both species were higher during summer and fall than during winter and spring. There was no significant difference in the catches of Humboldt sucker and white catfish between the two time periods. Tem- poral catch trends within the time periods are indicated by the significant differences in replicates for all species except the white cat- fish, which was scarce in catches during both periods. Environmental Variables Fish catch and limnological data were tab- ulated by sample period, station, and depth (Vigg 1979). Seasonal variations of environ- mental parameters were pronounced (Table 4). Temperature was highest during August (maximum 27.0 C), and lowest in late De- cember (1.0 C). The surface DO concentra- tion was never below 8.0 mg/1. Bottom DO deficits occurred during August and Septem- ber. During early August, when maximum annual water temperatures occurred in the upper end of the reservoir, bottom DO de- creased there to 2 mg/1. Bottom DO was de- pleted in midreservoir in late August and in the lower end in September (Fig. 4). This DO depletion trend probably indicated either movement of the low-oxygen water mass down reservoir or differential in-place bot- tom DO depletion, or both. Destratification and mixing in the upper end of the reservoir in late August resulted in high DO concen- trations (10 mg/1) throughout the water col- umn. Destratification and reoxygenation of the midreservoir area took place during late September, and by October the entire reser- voir was well mixed. Dissolved oxygen con- centrations were near saturation levels for the rest of the year. Definite seasonal variation in turbidity oc- curred. Surface turbidity was highest (max- imum of 79 JTU during February at Station 4) during the winter and spring, when high rainfall, runoff, and erosion resulted in large amounts of suspended sediments in the lake. Inorganic suspended sediments persisted in the bottom zone of the lake throughout sum- mer. Turbidity was lowest during fall, when Table 2. Percentage fish catch (adjusted to 12-h set) by station and season in Ruth Reservoir, June 1974 through May 1975. Season and Species samples per Humboldt Golden Brown White Rainbow Largemouth station Station sucker shiner bullhead catfish trout bass Total Summer 7 1 17 7 < 1 11 38 0 10 7 2 8 7 1 23 0 0 6 7 3 13 6 < 1 23 13 18 8 7 4 32 26 32 26 25 18 30 7 5 31 54 66 17 25 64 47 FalP 7 1 21 8 6 41 11 25 15 7 2 19 2 10 33 11 25 13 7 3 19 31 3 7 33 0 18 7 4 41 60 81 19 45 50 55 Winter" 6 1 25 9 18 100 0 0 19 6 2 3 0 3 0 0 0 2 6 3 15 4 6 0 0 100 10 6 4 57 98 74 0 100 0 70 Spring 6 1 1 0 0 13 0 0 1 6 2 4 7 5 26 0 0 7 6 3 11 0 5 26 0 0 6 6 4 22 77 39 30 0 0 52 6 5 62 16 52 4 0 0 34 "Catches at Station 5 were excluded because this lake area was dry in fall and winter. December 1982 ViGG, Hassler: Ruth Reservoir Fish 535 all suspended sediments had been flushed from the reservoir. Turbidity was highest at the bottom and lowest at the surface during all seasons. Lake area effects also introduced considerable variation in turbidity— i.e., spa- tial trends in turbidity occvirred as storm nm- off moved through the reservoir. Conductivity varied with time and with vertical and horizontal lake area. However, variation was not great, the values ranging only from 80 to 200 jumho/cm (mean, about 125 jumho/cm). Simple and Multiple Linear Correlations Correlations between fish catch and con- current measurements of environmental vari- ables at specified stations indicated that tem- perature and turbidity had major effects on fish catches (Table 5). Consistent significant direct temperature and inverse turbidity rela- tionships with the catches of Humboldt suck- er, brown bullhead, white catfish, and total species occurred. Although a significant inverse relationship existed between fish catches and DO, there was no discernible biological basis for a cause-effect relationship of this type; i.e., in- creased DO concentrations would not be ex- pected to cause a decrease in fish catches. The range of DO saturation variation was not great, and DO concentrations measured at the water depths of net sets were not limiting to fish. Since there was generally a high cor- relation (r>0.90) between temperature and DO concentration, it is reasonable to assume that the fish-DO correlation is a result of the indirect temperature effect. Conductivity and fish catches were not consistently related. Environmental variables with biologically explainable effects on fish catch were in- cluded in multiple linear correlations with fish catch (Table 6). Significant (P<0.01) multiple linear correlations existed between total and individual species catch and the turbidity-temperature environmental system. Surface turbidity and bottom temperature accounted for 80.5 percent of the variation in total fish catch. Time of year, depth of sam- pling station, and Mad River inflow ex- plained very little additional variation in total fish catch. This pattern was consistent for all major fish species. Turbidity and tem- perature accounted for 72.2, 53.5, and 58.8 percent of the catch variation for Humboldt suckers, brown bullheads, and white catfish, respectively. A significant (P<0.05) relation- ship also existed between the turbidity-tem- perature system and the catch of golden shi- ners. However, the proportion of catch variation explained by temperature and tur- bidity—about 30 percent— was substantially less for the golden shiner than for the other species. In all tests, inclusion of additional en- vironmental variables did not account for a statistically significant proportion of inde- pendent variation. The catches of largemouth bass and rainbow trout were so small that correlation analyses would not be meaningful. Discussion The fish population dynamics of the reser- voir have not been continuously monitored since the dam was completed in 1961. How- ever, current evidence does suggest that es- tablishment of nonnative species in the late 1960s was associated with a decline in the rainbow trout population. Introductions of the golden shiner, brown bullhead, white cat- fish, and largemouth bass were unauthorized. Golden shiners were first observed during Table 3. Gill net catches (adjusted to 12-h set) at Sta- tion 5 during summer-fall (s-f) and winter-spring (w-s) in Ruth Reservoir June 1974 through May 1975. Species, and seasonal period Number of Catch sets Total Mean Humboldt sucker s-f 10 223 22.3 w-s 9 171 19.0 Golden shiner s-f 10 429 42.9 w-s 9 45 5.0 Brown bullhead s-f 10 305 30.5 w-s 9 12 1.3 White catfish s-f 10 9 0.9 w-s 9 1 0.1 Total s-f 10 978 97.8 w-s 9 233 25.9 536 Great Basin Naturalist Vol. 42, No. 4 summer and fall 1968 (La Faunce 1968), and brown bullhead and white catfish during fall 1968. Largemouth bass are believed to have been introduced later— possibly in 1970. Changes in relative abundance of adult fish of different species are apparent from com- parisons of gill net samples taken in 1968 with those taken during the present study. Rainbow trout composed 31 percent of the catch in 1968, but less than 1 percent in 1974-1975. The relatively high trout catch in 1968 probably represents a population that remained from the stocking of hatchery- reared fish in the previous May and the resi- dent river population entrapped by the dam. Corresponding to the dramatic difference in trout catches were the substantial differences in catches of golden shiners (from 15 to 33 percent), brown bullheads (from 0 to 20 per- cent), and white catfish (from a trace to 3 percent). Humboldt suckers made up 54 per- cent of the catch in 1968 and 43 percent in 1974-1975. The largemouth bass maintains a naturally reproducing population and sup- ports a sizable fishery in the reservoir. It was probably more abundant in 1974-1975 than the gill net samples indicated (<1 percent) because centrarchids are typically difficult to capture in nets. Crayfish, which were very abundant in the 1968 sample, were present only in trace amounts in 1974-1975. Western suckers and golden shiners are two of the most successful competitors of rainbow trout in terms of reduced trout pro- duction in California reservoirs (Inland Fish- eries Branch 1971). Humboldt sucker and golden shiner composed over 75 percent of the sample in numbers and 83 percent in weight during 1974-1975. Thus, the 1974-1975 species composition and relative abundance of nongame fish in Ruth Reservoir could have been a factor detrimental to the reservoir trout population. Erman (1973) re- ported that populations of {Catostomus ta- hoensis) and (C. platijrhynchus) in Sagehen Creek increased from 17.8 percent (1952-1961) before impoundment to 41.3 percent in Stampede Reservoir and 79.2 per- cent upstream (1970-1972) after impound- ment. This illustrates that this stream-reser- voir system favored sucker populations. Since Humboldt suckers spawn in the Mad River during the same (spring) season as rain- bow trout, it is likely that the young of the two species compete for space and food. At present, the natural reproduction of trout in the Mad River above Ruth Reservoir appears negligible. The rainbow trout fishery of the reservoir has been maintained by stocking fingerling (1962-1968) and catchable-size fish (1969-1975). Since these hatchery-raised trout compete unsuccessfully with other res- ervoir species, the recent strategy of stocking catchable trout during times of heavy angler effort (i.e., before the opening weekend of the fishing season and before holidays) on a put-and-take basis is logical. However, if trout of a more predaceous strain were plan- ted at a larger size, they would be able to forage on golden shiners. Both temporal availability and spatial dis- tribution of the fish in Ruth Reservoir were associated with environmental properties that varied on a seasonal basis. A cyclic trend of high catches during the warm summer and fall, and low catches during the cold and rainy winter and spring was apparent for all species. The environmental-fish relationships quantified during this study were simple; i.e., temperature was directly related, and turbi- dity inversely, to fish catches. Both temper- Table 4. Mean seasonal environmental measurements at Im below the surfaces (S), mid-depth (M), and Im above the bottom (B) at Stations 1-4 in Ruth Reservoir, June 1974 through May 1975. Temperature (C) 1 S 'urbidit) (JTU) M 21.9 6.0 34.1 29.4 22.9 f B Conductivity (/xmho/cm at 25 C) S M B Dissolved oxygen (mg/1) Season s M B S M B Summer Fall Winter Spring Annual mean 22.2 16.5 4.6 8.6 13.0 18.3 16.1 4.4 7.8 11.7 14.8. 15.6 4.6 7.2 10.6 14.8 5.7 30.5 27.6 19.7 41.8 8.2 39.9 34.5 31.1 101.9 99.7 97.3 146.8 145.2 142.2 133.4 131.3 126.4 165.4 108.0 115.8 136.9 121.1 120.4 10.0 9.5 13.1 12.2 11.2 9.0 8.0 8.9 7.9 13.0 12.8 12.1 12.0 10.8 10.2 December 1982 ViGG, Hassler: Ruth Reservoir Fish Dissolved Oxygen 537 O a> o Q. E 25 20 Temperature s= Surface M=Middepth B= Bottom Station I. Station 2 Station 3. Station 4. 15 - 10 _ E c Q) o» X O a> _> o (O CO 0 25 20 15 10 0 August August September September 6 23 4 19 Fig. 4. Bottom-dissolved oxygen concentration and vertical temperature profile bv station in Ruth Reservoir Au- gust and September 1974. atiire and turbidity have known biological relationships affecting the survival, phys- iology, and behavior of fish. The mean summer temperature in Ruth Reservoir (22.2 C) approximates the pre- ferred temperature of the two most abundant fish species— Humboldt sucker and golden shiner (Reutter and Herdendorf 1974). The maximum surface temperature of Ruth Reser- voir (27 C) exceeds the upper lethal threshold 538 Great Basin Naturalist Vol. 42, No. 4 Table 5. Significant correlation coefficients between fish catch and selected environmental variables measured at surface (S), mid-depth (M), and Bottom (B) at Stations 1-4 in Ruth Reservoir from June 1974 through May 1975 (26 observations per station, n= 104). Temperature Turbidity Species S .213° ..381°° .269°° .411°° .343°° M B S M B Rainbow trout Largemouth bass Humboldt sucker Golden shiner Brown bullhead White catfish Total catch .272°° .436°° .214° .347°° .460°° .431°° .311°° .477°° .276°° .417°° .482°° .519°° -.214° -.383°° -.201° -.323°° -.350° ° -.271°° -.400°° -.233° -.215° -.360°° -.391°° -.202° -.369°° -.269°° -.243° -.289°° -.421°° °P<0.05 "P<0.01 of rainbow trout (National Academy of Sci- ences 1972); thus, the summer temperature regime restricts the spatial distribution of the trout. All other species, however, are capable of inhabiting the productive littoral and sur- face waters during the entire growing season. In fact, higher temperatures than those in Ruth Reservoir would favor largemouth bass and catfish, since their temperature for opti- mum growth exceeds 27 C (Strawn 1961, An- drews and Stickney 1972, Kilambi et al. 1970, Crawshaw 1975). The linear area sampling design provided a description of the fish species habitat prefer- ence in relation to seasonal environmental dynamics. The relatively shallow upper end provides most of Ruth Reservoir's littoral area, and the river-reservoir interface. It rep- resents an important habitat for the adult fish population, and probably serves as a spawn- ing and nursery area. Humboldt sucker and golden shiner primarily inhabited this region, regardless of season. Brown bullhead likewise preferred this habitat during all seasons; how- ever, the relative proportion of this species in the upper end of the reservoir increased dur- ing the summer. The relation between higher catches and the warm upper reservoir area indicates an interaction with temperature, since the upper end was the first to warm during spring and remained the warmest area of the lake throughout summer. Turbidity and the associated high inflow- outflow were inversely related to fish catch; the behavior and distribution of all fish spe- cies were negatively influenced by turbid wa- ter. Turbidity, inflow, outflow, flushing rate, and water level are all closely interrelated in Ruth Reservoir. Water level fluctuations af- fect the reproductive success of reservoir- spawning fish such as largemouth bass, white catfish, brown bullhead, and golden shiners in terms of spawning habitat, nursery habitat, and food availability. Drawdowns resulting in elimination of the shoals during the spawning season can cause direct mortality to eggs and larvae. Since larvae and juvenile fish feed on zooplankton, an abundant population of zoo- plankton is necessary at the time fish eggs hatch. Lider (1977) documented the deple- ting effect of the flushing rate on zooplank- ton populations in Ruth Reservoir; appre- ciable populations did not develop during spring until surface spilling ended. Young fish can also be lost in the discharge of a reser- voir. Walberg (1971) reported that the timing and rate of flushing affects the year-class sur- vival of reservoir fish. In addition to biological parameters, in- flow and outflow play a major role in the de- velopment of water quality patterns within a reservoir (Wunderlich 1971). Therefore, fac- tors that alter the flow regime— downstream water needs, modification of the dam, or modification of the outlet structure, would affect the biotic and abiotic ecosystem of Ruth Reservoir. Summary and Conclusions The fish population of Ruth Reservoir was dominated by Humboldt sucker, golden shin- er, and brown bullhead. White catfish formed an appreciable proportion of the population. Although largemouth bass were not heavily represented in the catch, their population is naturally maintained and their actual abun- dance is probably substantial. The species December 1982 ViGG, Hassler: Ruth Reservoir Fish 539 Table 5 continued. Conductivity Dissolved oxgyen Sampling period Sampling station S M B S M B .213° .214° .210° .231° .203° -.283°° -.393°° -.256°° -.261°° -.496°° -.436°° -.247° -.469°° -.198° -.195° -.453°° -.398°° -.376°° -.230° -.415°° -.320°° -.195° -.406°° -.248° -.223° .307°° .365°° .320°° .443°° composition and relative abimdance of the Ruth Reservoir fish population is not con- ducive to salmonid production. The rainbow trout population is negligible and maintained by put-and-take stocking. Catch of all fish species was positively correlated with tem- perature and negatively correlated with tur- bidity and inflow-outflow. Management practices affecting these environmental pa- rameters would thus effect the fish popu- lation of Ruth Reservoir. We believe the following measures would be beneficial in the management of the Ruth Reservoir fishery: 1. Promote the Ictalurid fishery, including night fishing. 2. Continue stocking catchable-size rain- bow trout during times of high angler effort. Study the feasibility of stocking more predacious salmonids (e.g., steel- head, Salmo gairdneri, or brown trout, Salmo triitta) at a larger size. 3. Study the detailed ecology of the large- mouth bass population. 4. Study the spawning potential for salmonids in the upper Mad River. 5. Undertake watershed management in the upper Mad River to alleviate soil erosion and the associated turbidity problems. Literature Cited Andrews, J. W., and R. R. Stickney. 1972. Interaction of feeding rates and environmental temperature on growth, food conversion, and body composi- tion of channel catfish. Amer. Fish. Soc, Trans. 101(l):94-99. California Department of Water Resources. Reser- voir limnology, Ruth Reservoir, 1969. Northern District. Red Bluff, California (unpublished data). CooLEY, W. W., and p. R. Lohnes. 1971. Multivariate Data Analysis. John Wilev and Sons, New York. 380 pp. Crawshaw, L. I. 1975. Attainment of the final thermal preferendum in brown bullheads acclimated to different temperatures. Coinp. Biochem. Physiol. A. Comp. Physiol. 52(1): 171-173. Erman, D. C. 1973. Upstream changes in fish popu- lations following impoundment of Sagehen Creek, California. Trans. Amer. Fish. Soc. 102(3) :626-629. Inland Fisheries Branch. 1971. Report on the status of reservoir research. California Dept. of Fish and Game. Inland Fish. Admin. Rep. 71-1. 45 pp. Iwatsubo, R. T., L. T. Britton, and R. C. Averett. 1972. Selected physical and chemical character- istics of 20 California lakes. U.S. Dept. Inter. Geol. Surv. Menlo Park, California. 59 pp. KiLAMBi, R. W., J. R. Noble, and C. E. Hoffman. 1970. Influence of temperature and photoperiod on growth, food consumption, and food conversion efficiency of channel catfish. Pages 519-531 in Proc. Southern Div. American Fish. Soc, Atlanta, Georgia. La Faunce, D. A. 1968. Gill net sampling at Ruth Lake, Trinity County. California Dept. of Fish and Game. Eureka, California (unpublished mimeo). 2 pp. Lider, E. L. 1977. The distribution and abundance of plankton and their association with environmen- tal variables in Ruth Reservoir, California. Un- published thesis. Humboldt State Univ., Areata, California. 88 pp. National Academy of Sciences. 1972. Water quality criteria 1972. U.S. Environ. Prot. Agency. 594 pp. Reutter, J. M., and C. E. Herdendorf. 1974. Labora- tory estimates of the seasonal final temperature preferenda of some Lake Erie fish. Proc. Conf. Great Lakes Res. (Int. Assoc. Great Lakes Res.) 17:59-67. SoKAL, R. R., and J. RoHLF. 1969. Biometry. W. H. Freeman and Co., San Francisco. 776 pp. Strawn, K. 1961. Growth of largemouth bass fry at vari- ous temperatures. Trans. Amer. Fish. Soc. 90:334-335. I 540 Great Basin Naturalist Vol. 42, No. 4 Table 6. Multi pl' s linear correlations betweer I fish catch and turbidity ; ind temperature at S Stations 1-4 in Ruth Reservoir from June 1974 through May 1975 (n = 26). Correlation Multiple correlation Ffor analysis Environmental coefficients coefficient of variance Species variables (r) i^ (R) .849°° R2 .727 onR Humboldt sucker Surface turbidity -.759°° .576 29.809°°° Bottom temperature .814°° .663 Brown bullhead Surface turbidity Bottom temperature -.579°° .728°° .335 .530 .731°° .535 13.228°°° White catfish Surface turbidity Bottom temperature -.586°° .766°° .343 .587 .767°° .588 16.439°°° Golden shiner Surface turbidity Discharge temperature -.547°° .450° .299 .203 .549° .301 4.963° Total Surface turbidity Bottom temperature -.835°° .833°° .697 .694 .897°°° .805 47.535°°° -P < 0.05 "P< 0.01 ■■'■°P< 0.001 U.S. Army Corps of Engineers. 1973. Draft environ- mental impact statement Butler Valley Dam and Blue Lake Project. San Francisco District, Corps of Engineers. 211 pp. ViGG, S. 1979. Distribution and relative abundance of fish in Ruth Reservoir, California, in relation to environmental parameters. Unpublished thesis. Humboldt State Univ., Areata, California. 70 pp. Walburg, C. H. 1971. Loss of young fish in reservoir discharge and year-class survival, Lewis and Clark Lake, Missouri River. Pages 441-448 in G. E. Hall, ed.. Reservoir fisheries and limnology. Spec. Publ. No. 8. Amer. Fish. Soc, Washington, D.C. WiNZLER AND Kelly. 1975. Ruth Lake Reservoir tur- bidity study. Winzler and Kelly, Environmental Consultants, Eureka, California. 27 pp. WuNDERLiCH, W. O. 1971. The dynamics of density stra- tified reservoirs. Pages 219-231 in G. E. Hall, ed.. Reservoir fisheries and limnology. Spec. Publ. No. 8. Amer. Fish. Soc, Washington, D.C. Young, J. C. 1971. Geology of the Mad River Basin. Pages 27-32 in Proceedings of the Mad River Symposium sponsored by the Office of Dean of Public Services and Department of Geology and Earth Science. Humboldt State College, Areata, California. TEMPERATURE AND SALINITY RELATIONSHIPS OF THE NEVADAN RELICT DACE Steven Vigg' Abstract.- The relict dace, Relictus solitarius, represents the only genus and species of fish native to Ruby Butte, Goshute, and Steptoe valleys in northeastern Nevada. In their natural habitats temperature ranges 0-25 C and salinity 175-1,158 nig/liter. The upper median thermal tolerance limit (96-hr TL50) of Butte Valley relict dace was 30.6 C when acclimated at 18-20 C. Relict dace tolerated total dissolved solids (TDS) of 11 043 mg/liter with no mortality during 96-hr exposures, but experienced 100 percent mortality at concentrations of 15 759 mg/liter with a mean resistance time of 23 hours. Tolerance of relict dace to 30 C was lowered as TDS was increased from 2 845 to 5,652 mg/hter. The Butte Valley fish were slightly more resistant to elevated salinities than the Goshute Valley sample, and conversely the Goshute Valley sample may be slightly more resistant to elevated temperature The relict dace, Relictus solitarius, is listed as "of special concern" by the Endangered Species Committee of the American Fisheries Society (Deacon et al. 1979). However, it was recently taken off the State of Nevada's pro- tected species list. Before the present study there were no quantitative data on the habitat requirements of relict dace. The purpose of this research was to determine a provisional upper lethal temperature limit (96-hr TL50), and to con- duct range finding tests on the total dissolved solids (TDS) tolerance of this unique fish. Relict dace apparently evolved during the past 1.5 to 2.0 million years in the contiguous drainage basins of pluvial lakes Franklin, Gale, Waring, and Steptoe just south of the conjoining parts of the Lahontan and Bonne- ville basins (Hubbs et al. 1974). As these Pleistocene lakes desiccated over the last 10,000 years, the relict dace was the only genus and species of fish to survive in the remnant springs of contemporary Ruby, Butte, Goshute, and Steptoe valleys, which comprise some 14,682 km2 in northeastern Nevada. The species also occurs in Spring Valley, where it is believed to be an in- troduction (Fig. 1). The taxon was first described by Hubbs and Miller (1972) from extensive research dating back to 1934. As well as being mor- phologically distinct from its closest relatives, Gila and Rhinichthys, this endemic cyprinid is also unique in terms of genetics (Lugaski 1980) and geographical distribution (Smith 1978). Relict dace was once the most abundant of the four fish species native to the north cen- tral Great Basin (Hubbs et al. 1974). The spe- cies is losing populations and habitat (Hardy 1979), due to predation and competition by exotic fish species, modification of natural springs into stock ponds and irrigation chan- nels, and groundwater mining. Methods Temperature and salinity bioassays were conducted on relict dace from November 1980 to March 1981. Test fish were trans- ported from their natural habitats in Ruby, Butte, and Goshute valleys (Fig. 1) to the Desert Research Institute (DRI) Bioresources Center laboratory in Reno. Upon arrival at DRI the fish were maintained in 425-liter holding tanks a minimum of four days for ob- servation of handling stress. During this time the fish were acclimated at 18-20 C in de- chlorinated Truckee River water prior to testing. Relict dace from three different sources (Franklin Lake, Atwood Ranch, and Phalan Spring) were tested for 96 hours over a tem- perature range of 29.0 to 34.3 C. Two rep- licates of five fish each were tested in static 19-liter aquaria for each treatment. The test 'Bioresources Center, Desert Research Institute. P.O. Box 60220, Reno, Nevada 89506. 541 542 Great Basin Naturalist Vol. 42, No. 4 O^eao" Idaho Elko Co. Nevada Wells <> Elko Co. Franwiln ike [Atwood Ranch ^Os Wendover o fPhalan Sprinj NVCurrie I /J ^ ^ f i>>^ \y f s / Ely ( White Pine C ]\ tone l^ous e N \ \ III :^ Fig. 1. Areal distribution of relict dace in five valleys in northeastern Nevada. The shaded area indicates known distribution of relict dace. Arrows mark the location of habitats where fish were collected. Asterisks (°) identify relict dace habitats documented by Hubbs et al. (1974). aquaria were placed in a thermostatically controlled water bath to maintain a constant temperature (±0.1 C). Five control fish were placed in dechlorinated Truckee River water in a 19-hter aquarium at room temper- ature (approximately 20 C) for each treat- ment; no control fish died. Table 1. Salinity related measurements of test media for TDS-alkalinity bioassays with relict dace, Relictiis solitarius. Total Specific conductance Test media TDS° (mg//) alkalinity (mg//) (jumhos/cm at 25 C) pH Truckee River 157 78 187 7.5 Vi Pyramid Lake 1/2 Distilled 2,845 639 4,730 9.2 IX Pyramid Lake 5,652 1,262 8,560 9.2 2X Pyramid Lake n,043 2,580 16,900 9.2 3X Pyramid Lake 15,759 3,670 22,900 9.2 4X Pyramid Lake 20,475 4,620 28,200 9.1 "Summation of all constituents. Test fish were not fed 24 hours prior to the bioassay nor during the tests. Dissolved oxy- gen (DO) and pH were measured at least once and usually three times daily. The aquaria were continuously aerated, and mean DO values always exceeded 6 mg/hter. Tem- perature was measured with a calibrated dig- ital thermometer at least three times daily in each aquarium. Range finding tests were conducted to de- termine the TDS-alkalinity tolerance of relict dace. Concentrated Pyramid Lake water (2X, 3X, 4X) was used to achieve elevated salinity levels (Table 1). Fish were derived from two sources, Atwood Ranch Spring and Phalan Spring, to test for differences by population. Procedures similar to those of the thermal bioassays were used: replicates of five fish/test in 19-liter continuously aerated aquaria with selected environmental param- eters monitored. Temperature was held con- stant (21.4-22.3 C) via the thermal water bath. Data were interpreted in terms of re- sistance time as well as mortality. December 1982 ViGG: Nevada Relict Dace 543 Another series of tests was performed to evaluate salinity-temperature interactions, utilizing only the Atwood Ranch Spring pop- ulation. Pyramid Lake water (IX), and V2 Pyramid - V2 distilled water were utilized as the test media for normal (~21-22 C) and elevated (~30 C) temperatures. For all salin- ity tests, Truckee River water (low TDS) was used for controls. Electrical conductivity and alkalinity determinations for each treatment were determined by the DRI water chem- istry lab, using standard analytical procedures (APHA et al. 1975). TDS was determined by summation of all measured constituents. The total and mean weight of each group of five test fish were determined (in water) to the nearest 0.1 gram prior to testing. Dead fish were immediately removed and pre- served in 10 percent formalin; time of each death was recorded. The criteria for death were no opercular movements and no re- sponse to touch. Statistical analyses were conducted utilizing programs from SPSS (Nie et al. 1975). Results Temperature Range finding tests were conducted on rel- ict dace from Franklin Lake in the Ruby Val- ley. No mortality occurred at 29.0 C during 96 hours, and 100 percent mortality occurred at 34.3 C within 27 hours (10 fish/test). More definitive results were derived from the Butte Valley (Atwood Ranch) population with four replicate tests over a range of 30-31 C (Table 2). The 96-hr median tolerance limit (TL50) for relict dace collected at the Atwood Ranch Spring was 30.6 C. Concurrent tests on relict dace from At- wood Ranch and Phalan Spring at 30.7 C in- dicated that the latter population may be slightly more resistant to high temperatures. The Phalan Spring population experienced only 40 percent mortality at 30.7 C, but the Atwood Ranch Spring fish suffered 80 per- cent mortality. The higher temperature toler- ance of the Phalan Spring population may be due to the long-term physiological acclimati- zation at 19.5-23 C compared to 16.5-21 C at the Atwood Ranch Spring (Table 3). TDS-Alkalinity Relict dace tolerated total alkalinity (HCO3 -I- C03=as CaCOa) and corresponding TDS of 2,580 mg/liter and 11,043 mg/liter, respectively, for 96 hours without experienc- ing any mortality (Table 4). TDS levels of 15,759 and 20,475 mg/liter resulted in 100 percent mortality of both populations of rel- ict dace tested. Analysis of variance illus- trated a significant difference in resistance times by concentrations (P< 0.001), fish pop- ulation (P<0.05), and interaction between "^f "'iL 'r^'"P^''a^"''e tolerance tests (96-hr) of Butte Valley relict dace (Relictus solitarius) conducted from No- vember 1980 to March 1981 in dechlorinated Truckee River water at mean acclimation temperatures of 18-20 C Sample size is five fish per test in replicate tests. Mean test temperature (C) 30.0 30.0 Control (21.6) 30.6 30.6 Control (22.7) 30.7 30.7 Control (21.3) 31.0 31.1 Control (21.7) Mean chemical characteristics Number of measurements 18 Fish Dissolved Mean Number Percent oxvgen weight dead mortality (mg//) pH (g) (96 hr) (10 fish) 6.4 8.75 1.82 0 6.5 8.20 1.46 0 0 7.3 8..50 2.14 0 6.8 8.76 2.08 2 6.8 8.17 2.40 3 50 7.6 7.98 1.62 0 6.5 8.54 2.24 4 6.6 8.36 2.28 4 80 7.6 8.15 2.08 0 6.1 8.39 1.68 5 6.1 8.51 3.20 5 100 7.3 8.03 1.70 0 544 Great Basin Naturalist Vol. 42, No. 4 the main effects (P< 0.001). An a-posteriori comparison showed that the Butte Valley (Atwood Ranch) fish were significantly more resistant to concentrated alkaline-saline wa- ters than the Goshute Valley (Phalan Spring) population (P<0.05). The resistance of time of Atwood Ranch fish was about twice that of the Phalan Spring fish at both the 15,759 and 20,475 mg/liter levels (Table 5). There exists an apparent interaction be- tween temperature and salinity tolerances of relict dace (Table 4). No mortality occurred at low temperatures (21-22 C) over a range of TDS from 157 to 5652 mg/liter. But at an elevated temperature (30 C) differential mor- tality occurred: 0, 20, and 60 percent at TDS levels of 157, 2845, and 5652 mg/liter, respectively. Discussion Temperature Compared to other families of freshwater fishes, the Cyprinidae generally occupy an intermediate rank of upper thermal toler- ance. Through acclimation, an upper lethal temperature exceeding 30 C is achieved by most cyprinids; at the highest tested acclima- tion levels eight species ranged from 29.3 Table 3. Environmental characteristics of selected re 1981. {Rhinichthijs atratiilus) to 38.6 C {Carrassius auratus) (Brett 1956). The upper lethal TL50 of 30.6 C herein determined for relict dace acclimated at 18-20 C falls within the range exhibited by other cyprinids. If the test fish were accli- mated to higher temperatures, it is likely that the upper lethal temperature would be sig- nificantly greater. The maximum water temperature I mea- sured in a relict dace habitat was only 23 C; however, Hubbs et al. (1974) recorded sum- mer temperatures as high as 25 C. Different populations of relict dace are subjected to a wide variety of temperature regimes. Non- thermal springs and shallow ponds may have great daily and seasonal changes with mini- mum temperatures near freezing (e.g., Frank- lin Lake). At the other end of the spectrum are thermal springs that never vary more than a few degrees in terms of diel, seasonal, and annual, presumably over extremely long time periods. For example, the head of Pha- lan Spring exhibited less than a 1 C diel change in February when air temperature ranged from 7.5 to -9.0 C, and seasonal wa- ter temperature variation was only about 3 C. In addition, there are various types of in- termediate gradations with respect to the temperature emanating from the springs and lict dace, Relichis solitaritis, habitats, April 1980-February Total dissolved solids (mg//) Total Temperature Summation of all Summation alkalinity Valley/ Site Date (°C) constituents (USGS 1979) (mg// CaCOa) Ruby Alkali Pond 12/80 2.0 1721 1157.8 912 Franklin Lake 12/80 .3.5 683.8 444.3 387 Steptoe Steptoe Ranch 12/80 20.8 388.5 255.6 215 Cardano Ranch 12/80 11.5 415 265.3 242 Goshute Phalan Spring 04/80° 07/80° 22 23 — 216 175 115 118 09/80° 22 - 190 132 Butte Atwood Ranch 04/80° 19 — 375 172 07/80° 21 — 325 182 09/80° 20 — 325 158 12/80 16.5 469.3 325.7 232 Odgers 04/80° 12 — 216 120 07/80° 20 — 205 115 09/80° 14 - 190 118 "Conducted by: Water Analysis and Consulting, Inc. 1980. Water Quality Study Wells Environmental Statement Area. Bureau of Land Management, Elko District. December 1982 Vigg: Nevada Relict Dace 545 the changes that occur in the length of streams and/ or depths of ponds. Limited data exist on thermal tolerance levels of cyprinid species inhabiting the Great Basin. Speckled dace {Rhinichthys os- culus) taken from intermittant Arizona streams at < 25 C had ultimate incipient up- per lethal levels of 33 C for juveniles and 32 C for adults (John 1964). The Borax Lake chub, Gila boraxobiiis, is endemic to a ther- mal lake in southeastern Oregon; the lake typically exhibits temperatures of 29-32 C, with extremes of 17-35 C (Williams and Wil- liams 1980). Another cyprinid from the northwestern Great Basin, the desert dace (Eremichthys acros), is endemic to thermal springs ranging from 18.5-40.5 C; this Ne- vada species tolerated temperatures of 2-37 C in the laboratory when acclimated at 23 G (Nyquist 1963). Virgin River spinedace {Le- pidomedu moUipinis), native to the Golorado River System, had a 14-hr upper lethal tem- perature of 31.2-31.4 G when acclimated at 20 G (Espinosa and Deacon 1978). In contrast, extensive research has been done on the thermal requirements of Gypr- inodontidae inhabiting the Death Valley re- gion of the southern Great Basin. Pupfish (Cyprinodon) acclimated at 10-20 G can tol- erate temperatures of 39-40 G (Brown and Feldmeth 1971, Otto and Gerking 1973, Feldmeth et al. 1974). Representatives of another cyprinodont genus (Crenichthys) are remnants of Pluvial river systems in the southeastern Great Basin. Crenichthys baileyi inhabits springs in the Moapa Valley at constant temperatures of 32.2 G (Kopec 1949). In the Moapa River C. baileyi and Moapa cariacea (Gyprinidae) oc- cupied habitats at temperatures of 27-32 G and 19.5-32 G, respectively (Deacon and Bradley 1972). Crenichthys baileyi and C. ne- vadae live in various spring outflows in the White River Valley and Railroad Valley, re- spectively, at temperatures ranging from 21 to 37.3 G (Hubbs et al. 1967). At Lockes Ranch spring complex C. nevadae were ob- served at temperatures of 18.3-37.8 G (Baugh and Brown 1980). Although individuals may survive for short periods of time at extremely high temper- atures, the maximum constant temperature occupied by a reproducing population of desert fish is rarely greater than 35 G (Soltz and Naiman 1978). This may be attributed to the differential tolerance of various life stages; e.g., Amargosa pupfish (Cyprinodon nevadensis) juveniles are most tolerant of ex- treme temperatures, with adults inter- mediate, and eggs least tolerant (Shrode 1975, Shrode and Gerking 1977). The repro- ductive tolerance range (<50 percent hatch) of C. nevadensis was 24-30 G or one-seventh the critical thermal tolerance range (Shrode Table 4. Salinity tolerance tests (96 hr) of relict dace Relictus solitarius, conducted during December 1980 and March 1981. The first series of bioassays tests interaction with temperature; the second series tests differences by fish population. Sample size is five fish per test. Population: Mean chemical characteristics Fish Number Mean TDS valley of Temperature DO. weight Number Percent (mg//) (habitat) measurements (C) (mg//) pH (g) dead mortality 157 Butte (Atwood) 9 21.6 7.3 8.50 2.14 0 (Control) " 30.0 6.4 8.75 1.82 0 0 " 30.0 6.5 8.20 1.46 0 2,845 Butte (Atwood) " 21.9 7.4 9.11 1.78 0 0 30.0 6.4 9.20 1.18 1 20 5,652 Butte (Atwood) " 20.9 7.3 9.21 1.60 0 0 29.9 21.4 6.4 7.6 9.20 9.24 2.22 3 60 11,043 Butte (Atwood) 12 2.14 0 0 Goshute (Phalan) 21.4 7.6 9.24 2.00 0 0 15,759 Butte (Atwood) 7 21.8 7.6 9.25 2.22 5 100 Goshute (Phalan) 4 22.0 7.4 9.13 1.92 5 100 20,475 Butte (Atwood) 2 22.3 7.1 9.09 1.90 5 100 Goshute (Phalan) 22.3 7.1 9.09 1.80 5 100 157 Butte (Atwood) 12 21.4 7.6 8.35 1.98 0 (Control) Goshute (Phalan) 21.4 7.6 8.30 6.02 0 0 546 Great Basin Naturalist Vol. 42, No. 4 and Gerking 1977). This example illustrates that multiple criteria are necessary to estab- lish the thermal requirements of a fish spe- cies. Thus, the TL50 determined in this study at a single acclimation regime is a useful baseline datum but is not comprehensive. TDS-Alkalinity The relict dace has experienced a variable environment during the past two million years in the Great Basin. Numerous cycles of Pluvial filling and interpluvial desiccation of large Pleistocene lakes occurred. Con- comitant changes in the salinity of the lacus- trine habitats undoubtedly took place. The variable content of TDS presumably has been a factor of considerable importance in the survival or extinction, and probably in the speciation of the populations of endemic fish- es (Hubbs et al. 1974). During the past 10,000 years the lacustrine habitats of the relict dace have completely dried up and the species persists in remnant springs. The vast reduc- tion of surface water to isolated springs is the outstanding feature of the habitats of native fishes in the Great Basin (Hubbs et al. 1974). The elevated salinity level that relict dace tolerated (11,043 mg/liter TDS) is over six times the maximum TDS level of any current relict dace habitat measured. The corre- sponding total alkalinity level (2,580 mg/liter) represents nearly three times that in any current habitats. The spring habitats are generally <250 mg/liter total alkalinity and <500 mg/hter TDS (Table 3); the most Table 5. Mean resistance time of Atwood Ranch (A) and Phalan Spring (P) relict dace, Relicttts solitarius, to highly alkaline-saline waters. Sample size is five fish per test. Mean resistance 95% confidence TDS time interval (mg//) Population (minutes) (minutes) 11,043 A Indefinite (>5760) ~ P Indefinite (>5760) - 15,759 A 1907 1204-2610 P 848 280-1416 20,475 A 213 121-305 P 111 70-152 saline habitats are ponds in the Ruby Valley, which exhibit total alkalinity levels as high as 912 mg/liter (TDS = 1,721 mg/liter). I hy- pothesize that relict dace evolved the phys- iological mechanisms to withstand highly sa- line-alkaline waters during the desiccation of their lacustrine environment. In contrast to relict dace habitats, Cypr- inodont environments in the southern Great Basin can be extremely saline— as much as 4.5 times that of sea water, composed pre- dominantly of NaCl (Hunt et al. 1966). The composition of one saline habitat, Cottonball Marsh, was about 78 percent NaCl, with al- kalinity not listed as a constituent (LaBounty and Deacon 1972). During laboratory studies with the ionic composition approximating that of sea water, the Cottonball Marsh pup- fish {Cyprinodon milleri) survived at 88,000 mg/liter TDS, with a few individuals toler- ating 130,000 mg/liter for several weeks (Naiman et al. 1976). Numerous field obser- vations of various Cyprinodon species living at salinities greater than 90,000 mg/liter are documented in the literature (Barlow 1958, Deacon and Minckley 1974). Thus, relict dace can tolerate only a frac- tion of the TDS at which Cyprinodonts exist. This is probably a reflection of the differ- ences in their respective evolutionary envi- ronments. Precursors of the Cyprinodontidae had marine affinities (Smith 1981), and the family is apparently preadapted to harsh physical and chemical conditions; this en- abled the group to further evolve tolerance to extremes of salinity and temperature (Mill- er 1981). In contrast, Cyprinidae is a primary freshwater family (Miller 1958). However, the ionic composition of their present envi- ronments and test media may also be a criti- cal factor; i.e., the alkalinity component could affect the TDS tolerance of both taxa. Salinity bioassays on relict dace at normal and elevated temperatures demonstrated a synergistic effect. Brett (1960) points out that with the multiple role of temperature bring- ing increased attention to the problem of in- teraction, emphasis will shift away from the singular effects of temperature to synergistic effects within the overall characteristics of environments that permit survival of the spe- cies. In his work with the euryplastic Cypr- inodon macularius, Kinne (1960) showed that December 1982 Vigg: Nevada Relict Dace 547 the combination of temperature and salinity was of basic physiological importance; the ef- fects of a given temperature depend on the salinity and vice versa. Similarly, when deal- ing with the habitat requirements of relict dace, the synergistic relationships of salinity and temperature are important with respect to their physiology and ecology. Acknowledgments This research was sponsored by the Desert Research Institute (DRI) Project Assignment Committee. Dr. Clifford J. Murino, DRI president, was very supportive of the work. Consultation with Dr. James E. Deacon (Uni- versity of Nevada, Las Vegas) was helpful in initiating this research on relict dace. Dr. Robert R. Miller (University of Michigan, Arm Arbor) kindly reviewed my proposal and manuscript for accuracy of content. Mr. Da- vid L. Galat (Colorado Cooperative Fishery Research Unit) and Dr. Thomas P. Lugaski (University of Nevada, Reno) critically read the manuscript and made helpful suggestions. DRI Water Center Analytical Laboratory conducted water chemical analyses. Literature Cited APHA (American Public Health Association), American Water Works Association, and Water Pollution Control Federation. 1975. Standard methods for the examination of water and wastewater, 14th ed. New York, New York USA. 1193 pp. Barlow, G. W. 1958. High sahnity inortaHty of desert pupfish, Cijprinodon macularis. Copeia. 1958:231-232. Bauch, T. M., and B. G. Brown. 1980. Field observa- tions on the response of the Railroad Valley springfish {Crenichthys nevadae) to temperature. Great Basin Nat. 40(4):359-360. Brett, J. R. 1956. Some principles in the thermal re- quirements of fishes. Quart. Rev. Biol. 3 1(2): 75-87. 1960. Thermal requirements of fish— three dec- ades of study. Pages 110-117 in C. M. Tarzwell, ed.. Biological problems of water pollution. United States Department of Health, Education, and Welfare. Robert A. Taft Sanitary Engineer- ing Center, Cincinnati, Ohio, USA. Brown, J. H., and C. R. Feldmeth. 1971. Evolution in constant and fluctuating environments: thermal tolerances of desert pupfish (Cijprinodon). Evolu- tion. 25(2):390-398. Deacon, J. E., and W. G. Bradley. 1972. Ecological distribution of fishes of Moapa (Muddy) River in Clark County, Nevada. Trans. American Fish- eries Soc. 101(3):408-419. Deacon, J. E., G. Kobetich, J. D. Williams, and S. Contreras. 1979. Fishes of North America en- dangered, threatened, or of special concern. Fish- eries (Bethesda) 4(2):29-44. Deacon, J. E., and W. L. Minckley. 1974. Desert fish- es. Pages 385-487 in Desert Biology, Vol. 2. G. W. Brown, Jr., ed. Academic Press, New York New York, USA. Espinosa, F. a., and J. E. Deacon. 1978. Rearing bait fish in the desert southwest. State of Nevada. Fi- nal Report to Federal Aid in Commercial Fish- eries Research and Development Project No. 6-9- D. University of Nevada, Las Vegas. USA. 29 pp. Feldmeth, C. R., E. A. Stone, and J. H. Brown. 1974. An increased scope for thermal tolerance upon acclimating pupfish {Cijprinodon) to cycling tem- peratures. J. Comp. Physiol. 89:39-44. Hardy, T. 1979. The Inter-Basin Area Report- 1979. United States Department of the Interior and the State of Nevada. Pages 5-21 in Proceedings of the Desert Fishes Council. Vol. XI. Bishop, Cali- fornia, USA. HuBBS, C, R. C. Baird, and J. W. Gerald. 1967. Ef- fects of dissolved oxygen concentration and light intensity on activity cycles of fishes inhabiting warm springs. Am. Midi. Nat. 77(1):104-115. HuBBS, C. L., AND R. R. Miller. 1972. Diagnosis of new cyprinid fishes of isolated waters of the Great Ba- sin of western North America. Trans. San Diego Soc. Nat. Hist. 17(8): 101-106. HuBBS, C. L., R. R. Miller, and L. C. Hubbs. 1974. Hy- drographic history and relict fishes of the north central Great Basin. Mem. California Acad. Sci. Vol. 7. 259 pp. Hunt, C. B., T. W. Robinson, W. A. Bowles, and A. L. Washburn. 1966. Hydrologic basin, Death Val- ley, California. U.S. Geol. Surv. Prof. Paper No 494-B. John, K. R. 1964. Survival of fish in intermittent streams of the Chiricahua Mountains, Arizona. Ecology. 45:112-119. Kinne, O. 1960. Growth, food intake, and food con- version in a euryplastic fish exposed to different temperatures and salinities. Phvsiol. Zool. 33(4):288-317. Kopec, J. A. 1949. Ecology, breeding habits, and young stages of Crenichthys baileyi, a cvprinodont fish of Nevada. Copeia 1949(1):56-61. ' LaBounty, J. F., and J. E. Deacon. 1972. Cijprinodon miUeri, a new species of pupfish (family Cyprino- dontidae) from Death Vallev, California. Copeia 1972(4):769-780. Lugaski, T. P. 1980. Comparative chemotaxonomy of selected Great Basin native cyprinid fishes. Un- published dissertation. Univ. of Nevada, Reno USA. Miller, R. R. 1958. Origin and affinities of the fresh- water fish fauna of western North .\merica. Pages 187-222 in Carl L. Hubbs, ed., Zoogeographv. American Association for the .'Advancement of Science, Baltimore, Publ. No. 51. 1981. Coevolution of deserts and pupfishes (Genus Cyprinodon) in the American Southwest. Pages 39-94 in R. J. Naiman and D. L. Soltz, eds., Fishes in North American deserts. Wiley & Sons, New York, New York, USA. 548 Great Basin Naturalist Vol. 42, No. 4 Naiman, R. J., S. B. Gerking, and R. E. Stuart. 1976. Osmoregulation in the Death Valley pupfish Cyprinodon milleri (Pisces: Cyprinodontidae). Copeia 1976(4):807-810. NiE, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, AND D. A. Bent. 1975. Statistical package for the social sciences, 2d ed. McGraw-Hill, New York, New York, USA. Nyquist, D. 1963. The ecology of Eremichthijs acros, an endemic thermal species of cyprinid fish from northwestern Nevada. Unpublished thesis. Univ. of Nevada, Reno, USA. Otto, R. G., and S. D. Gerking. 1973. Heat tolerance of a Death Valley pupfish (Genus Cyprinodon). Physiol. Zool. 46(l):4,3-49. Shrode, J. B. 1975. Developmental temperature toler- ance of a Death Valley pupfish {Cyprinodon nevadensis). Physiol. Zool. 48:378-389. Shrode, J. B., and S. D. Gerking. 1977. Effects of con- stant and fluctuating temperatures on reproduc- tive performance of a desert pupfish, Cyprinodon n. nevadensis. Physiol. Zool. 50(1): 1-10. Smith, G. R. 1978. Biogeography of intermountain fi.sh- es. Intermountain biogeography: a symposium. Great Basin Nat. Mem. No"! 2:17-42. Smith, M. L. 1981. Late Cenozoic fishes in warm deserts of North America: a reinterpretation of desert ad- aptations. Pages 11-38 in R. J. Naimann and D. L. Soltz, eds.. Fishes in North American deserts. Wiley & Sons, New York, New York, USA. SoLTZ, D. L., and R. J. Naiman. 1978. The natural his- tory of native fishes in the Death Valley system. Natural History Museum of Los Angeles County, Science Series 30:1-76. USGS (United States Geological Survey). 1979. Tech- niques of water-resources investigations of the United States Geological Survey. Chapter Al. Methods for determination of inorganic sub- stances in water and fluvial sediments (M. W. Skougstad, M. J. Fishman, L. C. Friedman, D. E. Erdmann, and S. S. Duncan, eds.) Book 5, Labo- ratory Analysis. Water Analysis & Consulting, Inc. 1980. Water qual- ity study. Wells environmental statement area. Conducted for Bureau of Land Management, Elko District Office. 27 pp. Williams, J. E., and C. D. Williams. 1980. Feeding ecology of Gila boraxobius (Osteichthyes: Cypr- inidae) endemic to a thermal lake in southern Oregon. Great Basin Nat. 40(2):101-114. OBSERVATIONS ON WOUNDFIN SPAWNING AND GROWTH IN AN OUTDOOR EXPERIMENTAL STREAM Paul Greger' and James E. Deacon' Abstract.- The response of woundfin to different substrates and current speeds was investigated in an outdoor expenmental stream. Fisfi spawned in groups of 15-20 over 5-10 cm rock substrate in a .06 to .09 m/sec current at a depth of 10 cm. Eggs were adhesive on the undersides of the rocks. Fish spawned under these conditions grew to ap- proximately 55-60 mm TL in 5 months. The woundfin minnow, Phgopterus argen- tissimus Cope, is an endangered cyprinid presently occurring only in the Virgin River system of Nevada, Arizona, and Utah. Recent attempts at reestablishing this species in other areas of its original range in the Gila River Basin of Arizona have been unsuccess- ful. Almost 70 percent of the original wound- fin habitat in the Virgin River has been ren- dered unreliable during part of the year by irrigation diversions (Deacon 1979). A variety of development projects involving modifica- tion of present river flows pose additional threats to the continued survival of the woundfin. This study was undertaken to help identify habitat requirements for successful spawning. Such information is essential to an analysis of probable environmental impacts of various proposed development projects on the Virgin River as well as to the selection of areas within the former range of the species that might be suitable for reintroduction. Data on growth rates was taken to aid in an inter- pretation of a large data base on length- frequency of woundfin in the Virgin River. Methods and Materials Approximately 50 adult woundfin were collected from the Virgin River near the in- flow of Beaver Dam Wash at Littlefield, Ari- zona, on 26 April 1980 and transported to the experimental stream facility on the Uni- versity of Nevada campus in Las Vegas, Ne- vada. Many of the adults were robust, ap- pearing in prime reproductive condition. Fish were immediately transferred to the stream facility upon arrival. The stream (Fig. 1) consists of two raceway sections, a narrow upper section and a wider lower section, both constructed of concrete. Two pools, 2 X 2 m with a maximum depth of 1 m connect the raceways. The upper raceway is approximately 7 m in length and 45 cm in width, with a maximum depth of 15 cm. The lower raceway is 7 m in length and 90 cm in width, with a maximum depth of 45 cm. Current is generated by a 115 volt, 6.7 amp centrifugal pump that draws water out from the west pool through a circular rock filter. Water is pumped into the top end of the upper raceway, flows into the east pool, and then through the lower raceway into the west pool. No filters were used to remove waste prod- ucts but fresh well water was added daily to replace seepage and evaporation. This, in combination with low fish densities, seemed sufficient to prevent accumulation of am- monia or other harmful substances. The sys- tem design provides a choice of current speeds in the upper raceway as indicated for an average water level in Figure 1. Current speed in the lower raceway was too slow to measure with a Marsh McBirney hand-held, paramagnetic current meter. We provided a choice of substrates by placing large rock (15-25 cm diameter), small rock (5-10 cm diameter), gravel, and sand in distinct segments in both raceways (Fig. 1). Daily observations of woundfin dis- 'Department of Biological Sciences, University of Nevada. Las Vegas, Nevada 89154. 549 550 Great Basin Naturalist Vol. 42, No. 4 Current Profile at 30CM intervals M./Sec. East Pool Spawning Area .06 N Key: 15-25 CM Large Rock Shelf I Fig. 1. Outdoor experimental stream at the University of Nevada, Las Vegas. 5-10 CM Rock 2-3 CM Gravel Sand ffl tribution and behavior in the stream system Ryan thermograph. Fish were collected, pre- were made. Water temperature was mea- served, and measured at intervals to docu- sured with a mercury thermometer and a ment growth rates of fry and fingerlings. i December 1982 Greger, Deacon: Woundfin Spawning and Growth 551 Collections are on deposit in the Museum of Natural History at UNLV. Results and Discussion Fish were placed in the experimental stream on 26 April 1980 and observed at ir- regular times each day. They usually occu- pied the deeper waters in the pools and swam in loosely organized schools. On 3 May a tightly packed wedge-shaped school of 15-20 adults was observed making many quick turns over the small rock (5-10 cm) substrate of the upper raceway where the current speed was .06-.09 m/sec and depth was 7-10 cm. The distinctive movements of this group were essentially confined to this 45 cm section and continued for about 40 minutes. Similar movements were noted on occasions over the next 9 days. These sub- sequent occasions involved 4-7 fish and lasted only 4-5 minutes. Water temperature was 25 C during each spawning event and fluctuated from 19 to 26 C during the period of spawning. Spawning movements involved quick movements of the tightly packed school over the small rock substrate. Occasional flashing occurred, and at irregular intervals one or a few fish would leave the group and swim downstream to the pool below. Similarly at irregular intervals one or a few fish would swim upstream from the pool below to join the group. When fish entered the school, the group frequently would become more com- pact and appear to stop and vibrate as a group around the new arrival. This was as- sumed to be the spawning act. Immediately following it, the fish-to-fish distance would increase slightly, some individuals would pick at the substrate, and the main group would continue its tight circling movements over the small rock substrate. Although sex was not determined, we presumed that mature males dominated the school and that the fish entering and leaving were largely mature females. Depth was uniform throughout the upper raceway. Current speeds of .06-. 09 m/sec oc- curred over small rock, gravel, and sand sub- strates. Spawning activity was confined to the small rock substrate, where adhesive eggs were dropped between the interstices of the 5-10 cm rock. Woundfin fry were first observed in the stream on 14 May. All fry were located in the lower section of the stream where currents were minimal (.01-.03 m/sec). Woundfin fry appeared to orient close to the edges of the raceway where current speed was slower or over a shallow shelf in one end of a pool. They did not occur over the deep water. The fry, when first observed on 14 May, were about 8-10 mm TL. Water temperature was about 20 C. Close examination of the presumed spawn- ing site on 15 and 17 May revealed adhesive eggs attached to the underside of the 5-10 cm rock in the upper raceway. This was fur- ther evidence that woundfin utilized the small rock substrate for spawning. Eggs were approximately 1.5 mm in diameter and had a brown center with a clear outer region. Some eggs were white and opaque. If a spawn did occur on 3 May 1980, the period of devel- opment is about 10-11 days. Data for other cyprinids is similar. For example, Rhinchthys oscidus in the Chiricahua Mountains in Ari- zona required 6 days for hatching at 18 C plus 7-8 days to swim up from the interstices of the gravel (John 1963). Fry of 9 mm total length were first ob- served to orient in current (.06 m/sec) over sand in the upper raceway on 15 May. These fry held position close to the bottom and edges of the raceway in slower currents. Fry also apparently selected areas of shade in the late afternoon. On 18 May woundfin fry ap- peared to be increasing their range of move- ments to include rock and sand substrates in the upper raceway. From 22 May to 12 June fry (10-13 mm) moved further upstream over gravel and rock substrates until by 24 June, 18.5 mm fish were observed moving over all substrates in the upper raceway. As woundfin increase in size, they develop an increasing ability to move into swifter waters. This sug- gests that in the environment woundfin of 18-20 mm total length are able to move away from areas with slow current into open water areas where current is moderate. Field observations are consistent with these data. On 4 July the first schooling activity of fin- gerling was noted in the east pool. Some chasing behavior of fingerlings by adult woundfin was also observed. Finally, on 8 July, the stream was partially drained and 552 Great Basin Naturalist Vol. 42, No. 4 about 100 woundfin fingerling were removed and transported to the National Endangered Species Hatchery at Dexter, New Mexico. Twenty-five woundfin were retained for fur- ther study of growth. Over the previous month, mortality related to an Ichthyoph- thirius infection had been substantial. Information on growth of woundfin is giv- en in Table 1. The first three weeks of the ex- perimental period show little or no change in mean size. In fact, week 3 shows a slightly lower mean length than the previous week. This could have resulted from a second spawning or it could signal a difficult transi- tion from yolk-dependent growth to food-de- pendent growth. Weeks 4 and 5 show a growth increment of about 2 mm per week. Weeks 5, 6, and 7 show approximate growth increments of 4 and 5 mm per week. Thereafter, monthly measurements (July, August, and September) show growth changes of 12.4, 15.1, and 5.6 mm, respectively. Woundfin hatched in early May had reached a mean length of 35.7 mm by the end of July, 50.8 mm by the end of August, and 56.4 mm by 25 September. Ob- servations were terminated on 25 September. The stream was colonized by aquatic plants {Chara sp. and Naja sp.) and by a vari- ety of macroinvertebrates (Ostracoda, Chydoridae, Chironomidae, Ephemeroptera, Anisoptera, and Coleoptera). Although a high protein commercial feed was supplied, it ap- peared that woundfin fry fed primarily on the natural foods present. These included Chironomid adults, pupae, and larvae and Table 1. Weekly growth of woundfin fry and finger- ling in an outdoor experimental stream. The mean length represents the average of all measurements made during the weekly period. X Length Range Week Date N (mm) (mm) 1 14-20 May 31 9.4 8.5-11.5 2 21-27 May 51 9.9 7.5-12.0 3 29 May-4 June 15 9.2 7.0-11.0 4 12 June 7 12.7 9.0-15.0 5 15-17 June 9 14.3 12.0-16.0 6 25 June 6 18.5 17.0-20.0 7 IJuly 7 23.3 21.0-25.0 11 27 July 14 35.7 32.0-39.0 14 30 August 7 50.8 48 -55.0 17 25 September 8 56.4 49 -61.0 Chydorid crustaceans. Fry appeared to be in good condition. Conclusions 1. A small rock (5-10 cm diameter) sub- strate was specifically selected as the spawn- ing substrate. 2. Spawning was accomplished when a gravid female joined a group of ripe males exhibiting spawning behavior over the prop- er substrate (depth was about 7-10 cm). 3. Woundfin eggs are adhesive and drop between the interstices of rocks, where they adhere to the underside. 4. Spawning of woundfin occurs primarily in the spring when water temperatures reach about 25-26 C during the diurnal temper- ature cycle. 5. Woundfin of 10-12 mm TL are re- stricted in their movements by current, but woundfin of 17.0-20 mm TL are able to move freely throughout the stream in a vari- ety of current speeds. 6. Woundfin grow to about 50 mm in four months, a mean growth rate of about three mm/week. Growth may then slow, but an av- erage size of 55-60 mm during the first growing season is not unlikely. Acknowledgments Thom Hardy assisted in collection, trans- portation, and observation of woundfin. Tina Hardy assisted in maintenance of the stream and with observations of the fish. Their en- thusiastic cooperation is much appreciated. This study was supported in part by Contract 14-16-0002-78-919 from the U.S. Fish and Wildlife Service. Permits were provided by the Arizona, Nevada, and Utah Fish and Game departments and by the U.S. Fish and Wildlife Service. Literature Cited Deacon, J. E. 1979. Endangered and threatened fishes of the west. Great Basin Nat. Mem. No. 3:41-64. John, K. R. 1963. The effect of torrential rains on the re- productive cycle of Rhinichthys oscuhis in the Chiricahua Mountains, Arizona. Copeia 1963(2):286-291. EARLY DEVELOPMENT OF THE RAZORBACK SUCKER, XYRA UCHEN TEXANUS (ABBOTT) W. L. Mincklev' and Eric S. Gustafson' ,\bstract.- Fertilized ova of razorback sucker, Xyraiichen texanus, were adhesive for 3 to 4 hours after fertiliza- tion. Cleavage was completed at 24 hours, gastrulation occurred at 34 hours, and blood circulation was established at 11/ hours. Hatching occurred from 5.2 to 5.5 days after fertilization. Larvae were from 6.8 to 7.3 mm TL at hatch- ing. Yolk was assimilated at 13 davs (10 mm TL). All fins were formed and had ossified rays at 64 days (27 mm TL) The unique nuchal keel appeared about 200 davs after fertilization. The razorback sucker, Xyraiichen texanus (Abbott), is endemic to the Colorado River basin. As with much of the southwestern ich- thyofauna (Pister 1981), it is dechning in abundance (Minckley 1983). A program was commenced in 1974 to develop means of propagating the species (Toney 1974) and to delineate certain aspects of its life history. We studied embryological, larval, and juve- nile development of the species in 1974-75. Although our data are somewhat outdated in light of recent studies of catostomid larvae (reviewed by Fuiman and Witman 1979), al- most nothing has appeared on the early life history of this unique species. Winn and Miller (1954) presented a key to postlarval fishes of the lower Colorado River basin that included photographs and some descriptions of young X. texanus. A photograph by Douglas (1952: Fig. 3) was reidentified by Winn and Miller as speckled dace {Rhi- nichthys oscuhis [Girard]) rather than X. tex- anus. The present paper thus describes and figures early life-history stages of the razor- back sucker as determined from hatchery-and laboratory-reared individuals. Methods Initial information on hatchery propaga- tion and rearing of razorback suckers origi- nated from adult fish seined near Cotton- wood Cove in Lake Mohave, Arizona- Nevada, in winter 1974, and was compiled by personnel at Willow Beach National Fish Hatchery (in part, Toney 1974). Eggs were stripped from females and fertilized, and de- veloping young were initially housed in an indoor raceway at a mean water temperature of 14 C, then transferred 13 days post- hatching to an outdoor raceway where water temperature averaged 15 C. Samples were preserved daily in 10 percent formalin during the first month, and intermittently thereafter. Late postlarval and juvenile phases described below are based on the 1974 cohort. Additional adults were trammel netted from below Hoover Dam and in the vicinity of Carp Cove in Lake Mohave in March- April 1975. Most males were in active spawning condition, but females were either spent or not yet mature. Suitable females were interperitonealy injected with human chorionic gonadotropin, which induced oocyte maturation. A few hours after in- jection about 5,000 eggs were stripped from a single female and immediately fertilized with sperm of two males, as has been ob- served in nature (Douglas 1952). It is notable that water-hardened eggs obtained from naturally-matured females in 1974 were 2.9 mm diameter, but comparable ova obtained from hormone-induced maturation were 1.8 mm diameter. This disparity is far greater than has before been recorded in catostomid ova diameters (Fuiman and Trojnar 1980). We assume it resulted from precipitous mat- uration after hormone injection, but have no 'Department of Zoology, .\rizona State University, Tempe, Arizona 85287. 553 554 Great Basin Naturalist Vol. 42, No. 4 further explanation. By the time of hatching, both sets of embryos were of comparable length. Developing embryos were maintained in an indoor raceway at Willow Beach for 28 hours at temperatures ranging from 13 to 17 C. Eggs were then transported to aquaria at Arizona State University, where development continued at a constant temperature of 20 C. Observations and measurements were re- corded from living specimens by a stereo- microscope equipped with ocular microme- ter and camera lucida, and with a range of magnification of 2 to 2,000 X. All measure- ments are of total length. Observations on the 1975 cohort were made hourly through the first 30 hours, then every 2 hours until after hatching. Illustra- tions of selected stages through prolarval de- velopment were prepared through use of a camera lucida and from photographs. Ovum through prolarval development presented here is thus based on the 1975 fish. Speci- mens were preserved periodically in aceto- formalic acid (AFA; 9.0 parts ethanol, 0.4 parts 40 percent formaldehyde, and 0.5 parts glacial acetic acid). When larvae began to swim and feed actively, samples were pre- served less frequently. Development of the razorback sucker was divided into four major phases as defined by Hubbs (1943): (1) embryological devel- opment, fertilization of the egg until hatch- ing; (2) prolarval development, hatching to absorption of yolk; (3) postlarval devel- opment, yolk absorption to ossification of pelvic fin-rays; and, (4) juvenile devel- opment, pelvic ray ossification to maturation of gonads. Development staging followed Ba- linsky's (1948) general pattern for cyprinid fishes. Descriptive terminology was derived from Ryder (1885), Stewart (1926), Tavolga (1949), Winn and Miller (1954), and Long and Ballard (1976). Results Embryological Development Stage 1: unfertilized egg; day 0, 0 hour, 1.5 mm diameter. Ova milky white and translucent. Stage 2: fertihzed egg; day 1, 1 hour, 1.8 mm diameter. Chorion transparent and yolk milky-white and translucent; animal pole not yet visible to unaided eye. Water-hardened eggs with greater specific gravity than water, ova demersal, chorion adhesive, ova adhering to substrate and one another. Stage 3: 2 blastomeres, 3 hours (Fig. lA); beginning of cleavage. Blastomeres trans- parent, approximately 0.5 mm total diame- ter. Animal and vegetal poles distinguishable to unaided eye; ova telolecithal, cleavage meroblastic. Ova no longer adhesive. AFA preservation causes animal cells to whiten and become opaque; yolk becomes yellow white; chorion remains transparent. Stage 4: 4 blastomeres, 5 hours; second cleavage. Blastomeres approximately 1.0 mm total diameter. Stage 5: 8 blastomeres, 6 hours; third cleavage. Blastomeres occupy 1.2- by 0.8-mm rectangle on animal pole. Stage 6: 16 blastomeres, 7 hours; fourth cleavage. Stage 7: 32 blastomeres, 9 hours; fifth cleavage (Fig. IB). Stage 8: 64 blastomeres, 10 hours; sixth cleavage. 128 blastomeres, 11 hours; seventh cleavage. Large-celled blastula (morula); no blastocoel. Blastomeres occupy 25 percent of yolk surface. Stage 9: small-celled blastula (morula), 14 hours. Individual cells distinguishable; blasto- meres bulging upward from round yolk mass, occupying 25 percent of yolk surface. Blasto- mere layers progressively thinner toward pe- riphery of blastoderm; no blastocoel. Stage 10: morula, 24 hours (Fig. IC). Indi- vidual cells indistinguishable except with high power and chorion removed; blastoderm with granular appearance; undersurface flat, lying on flattened surface of yolk sphere; no blastocoel. Cleavage terminated. Stage 11: blastula (epiboly of blastoderm), day 2, 28 hours; 1.8 mm diameter (Fig. ID). Blastoderm spreading over yolk sphere and thinning (blastodisc). Blastocoel present. Peri- blast visible as cellular rim along periphery of blastoderm, beginning formation of inner layer of yolk sac. Outer layer of yolk sac to be formed from epiblast derived from blasto- derm. Blastoderm no longer bulging from yolk, capping under 33 percent of sphere. December 1982 Minckley, Gustafson: Razorback Sucker Development 555 Stage 12: early gastrula, 34 hours (Fig. IE). Underrim of blastodisc thickened to form "randwtibt" or marginal ridge with inner layer termed the germ ring. Ring thickest posteriorly, recognized as embryonic shield. Presumptive endodermal cells at posterior edge of shield beginning to involute through blastopore and spread beneath blastoderm. Cells of prechordal plate and notochord mi- grating inward over dorsal lip of blastopore (establishment of embryonic axis). Presump- tive mesodermal cells also turning inward, positioned either side of embryonic axis be- neath ectoderm and above endoderm. Peri- blast, randwulst cells, and germ ring cells not involved in involution spread over 50 percent of yolk sphere. Stage 13: middle gastrula, 35 hours. In- vagination lengthening embryonic shield; blastopore marks posterior axis of embryo. Stage 14: late gastrula, 36 hours (Fig. IF). Embryonic shield nearly reaching animal pole of egg on dorsal meridian; shield ap- proximately 1.3 mm long; concentration of invaginated cells clearly visible at anterior end of shield; randwulst cells and germ ring cells, accompanied by presumptive ectoder- mal cells, forming outwardly as epiblast; marginal ridge shifted below equator of egg; uncovered portion of yolk protrudes as yolk plug. Stage 15: early neurula, 45 hours, 1.8 mm diameter. Blastopore closed; yolk plug no longer protruding. Embryonic shield approx- imately 2.0 mm long, circumscribing 75 per- cent of egg, overlying yolk sac. Neural plate formed, lateral and anterior margins not clearly delimited. Notochord in form of ridge pressing into yolk, lateral notochord rudi- ment not clearly separated from sheets of mesoderm. Stage 16: late neurula, 47 hours (Fig. IG). Neural keel remains 2.0 mm long, circum- scribing 75 percent of egg. Neural plate con- tracted, more defined along edges. Cephalic region arrow shaped in dorsal view; eye rudi- ments forming; neural ridges as folds on ei- ther side of neural plate. Notochord sepa- rated from mesoderm. Stage 17: eye rudiments, day 3, 49 hours, embryo length 2.0 mm. Brain cavities and spinal cord formed by contraction of neural plate. Eye rudiments as lateral protrusions at G ^ Late Neurula t r- -— -\np \\ 1 ' \ -^ -^ ^YS Dorsal j—-^ Lj Oval^^eye & rf A Mes 1 Dorsal H ^___^Rh V/M L Jap "'Ml/^ ~~-~p^LPM ■Som Fig. 1. Early embryological stages of razorback suck- er, Xyrauchen texanus; chorion removed in all but A. Legend: AIP = anterior intestinal portal, Ant. = ante- rior, AP = auditory placode, Bl = blastoderm, Blc = blastocoel, Elm = blastomere, Bp = blastopore, Ch = chorion, ES = embryonic shield, GR = germ ring, HF = head fold, LPM = lateral plate mesoderm, Mes = mesencephalon, MR = marginal ridge, NF = neural fold, NK = neural keel, NP = neural plate, OP = optic placode, Pb = periblast, PM = perivitelline membrane, PS = perivitelline space, Rh = rhombencephalon, som = somites, Y = yolk plug, and YS = yolk sac. anterior end of brain. Prosencephalon (fore- brain), mesencephalon (midbrain) and rhomb- encephalon (hindbrain) distinguishable. Audi- 556 Great Basin Naturalist Vol. 42, No. 4 ^arl V Tai I bud Fig. 2. Early (A) and late (B) tailbud embryological stages of razorback sucker, Xyrauchen texanus; chorion re- moved-embryo extended in "B." Legend as in Figure 1 when applicable, and AV = auditory vesicle, Di = dien- cephalon, FG = foregut, L = lens, Met = metencephalon, Myl = myelencephalon, N = notochord, OC = optic cup, OV = optic vesicle, TB = tailbud, and Te = telencephalon. tory vesicles at level of hindbrain; first pair of somites present. Stage 18: cavities in eye rudiments, 53 hours. Optic placodes elongated, containing narrow vesicles; head-fold visible; sub- cephalic pocket and anterior intestinal portal ventral to head. Auditory placodes formed; 14-16 pairs of somites. Stage 19: oval eyes, 57 hours, 2.5 mm (Fig. IH). Optic placodes rounded to oval; vesicles not yet present in auditory placodes; 30 pairs of somites. Tail process distinguishable. Stage 20: early tailbud, 65 hours, 3.0 mm (Fig. 2A). Optic vesicles flattened on outer margins; lens forming. Auditory placodes with small vesicles. Tailbud developed, pro- truding from yolk sphere. Foregut forming. Stage 21: late tailbud, day 4, 78 hours, 3.8 mm (Fig. 2B). Anterior portion of embryo (head and anterior trunk) overlying yolk sphere; posterior portion (posterior trunk and tail) overlying cylindrical yolk mass. Optic stalks, cups, and lenses distinguishable. Pros- encephalon divided into telencephalon (fu- ture cerebrum) and diencephalon (future epithalamus, thalamus, and hypothalamus); rhombencephalon divided into meten- cephalon (future cerebellum) and myelen- cephalon (medulla oblongata). Heart rudi- ment present; tailbud lengthening, embryo motile within chorion. Stage 22: heart beat, 83 hours, 4.0 mm. Heart pulsations noted. Tail at right angle to body axis. Kidney ducts (pronephric ducts) and dorsal aorta formed ventral to neural tube and notochord; alimentary tract lined by endoderm and nearly complete; stoma- deum and proctodeum not apparent. Stage 23: fin fold, day 5, 103 hours, 5.3 mm. Fin fold appearing on tail and posterior dorsum; head growing outward from yolk sac. Circulatory system formed; nasal pla- codes present; tail beginning to straighten. Stage 24: blood circulation, 117 hours, 6.8 mm (Fig. 3). Head extending from yolk. Heart visible within pericardial cavity; flexed, sinus venosus and atrium lying above and left of ventricle and conus arteriosis; endocardium and epimyocardium dis- tinguishable. Three visceral (gill) arches formed; pupils visible within eyes, brown pigment granules in choroid regions. Yolk re- duced to cylinder below body axis. Blood flow: atrium -* ventricle -* conus arteriosus ^ ventral aorta -* branchial afferents ^ bran- chial efferents ^ dorsal aorta and internal carotid arteries -* vitelline artery -* caudal vein -> posterior cardinal vein -* anterior car- dinal and vitelline veins -* common cardinal vein (Duct of Cuvier) ^ sinus venosus -* atrium. Embryos extremely motile, some be- ginning to rupture chorion. Stage 25: pectoral fin rudiments, 120 hours, 6.8 mm. Pectoral fin anlagen appear- ing posteriad and ventrad to auditory vesicles. Tail slightly upturned. Stomadeum December 1982 Minckley, Gustafson: Razorback Sucker Development 557 Fig. 3. Embryo of razorback sucker, Xijrauchen texanus, at initial circulation of blood; embrvo extended and cho- rion not depicted. Legend as in Figures 1 and 2 when applicable, and ACV = anterior cardinal vein At = atrium CA = carotid artery, CCV = common cardinal vein (duct of Cuvier), CdV = caudal vein, Co = conus artriosus' DA = dorsal artery, FF = fin fold. My = myomeres, NP = nasal placode, P = pupil, PCV = posterior cardinal vein, SV = sinus venosus, VA = vitelline artery, Vt = ventricle, and VV = ventral vein. clearly visible. Myotomes discernible above yolk. Embryos flex once every 12 to 13 heart beats. Stage 26: pectoral fin buds, day 6, 125 hours, 7.3 mm. Rudiments of pectoral fins protrude, slightly flattened dorso-ventrally. Head continuing to extend outward from yolk. Prolarval Development Stage 27: hatching, 131 hours, 7.3 mm (Fig. 4). Pectoral fin buds paddle shaped. Tail straightened and median fin fold developed anteriorly on dorsum. Head flexed 45 degrees relative to body axis. Proctodeum discernible; eye pigment increased. Embryos scarcely motile, flexing along bottom with no directed movements. Stage 28: 142 hours, 7.5 mm. Four gill arches visible; head remains at 45 degree angle to body axis. Pectoral fln buds thin and broadly rounded; ventral fln fold appearing behind proctodeum. Stage 29: 144 hours, 7.5 mm. Lower jaw formed, not yet reaching level of eye; mouth oriface round. Head angle less than 45 de- grees relative to body axis. Stage 30: early prolarva, day 7, 162 hours, 8.0 mm (Fig. 5). Head straightened. Lower jaw reaching midlevel of eye, not movable. Pectoral fin differentiated into muscular lobe and membrane, not movable. Ventral fin fold developed to most anterior extension, em- bryonic fin membrane (continuous median fin) complete. Opercular membranes form- ing. Optic pigmentation brown, granular, al- most complete; no other melanophores. Stage 31: middle prolarva, day 9, 238 hours, 9.0 mm. Rudimentary gas bladder evi- dent. Lower jaw to anterior border of eye, movable; mouth rounded and terminal. Opercle covers anteriormost gill. Pectoral fin 0.5 mm long, movable. Proctodeum yet Fig. 4. Prolarva of razorback sucker, Xyrauchcn texanus. at hatching; legend as in Figures 1 to 3 when applicable and AT = alimentary tract, CFf = caudal fin-fold, DFf = dorsal fin-fold, GA = gill arches, PC = pericardial cav- ity, PFb = pectoral fin-bud, Pr = proctodeum, St = stomadeum, and VFf = ventral fin-fold. 558 Great Basin Naturalist Vol. 42, No. 4 Fig. 5. Early prolarva of razorback sucker, Xyraitchen texanus; legend as in Figures 1 to 4 when applicable, and Eph = esophagus, OpF = opercular flap, Pfmbr = pectoral fin membrane, and PFms = pectoral fin musculature. closed, yolk much reduced. Eye pigmentation completed, becoming black. Brown, stellate melanophores on ectoderm overlying mid- and hindbrain, and on paired dorsal pigment line, paired dorso-visceral pigment line (up- per body cavity), and unpaired midventral pigment line. No melanophores on horizontal myoseptum. Larvae swim to surface and feed on ground aquarium fishfood (Tetramin"^). Resting heart rate 120 beats per minute. Stage 32: late prolarva, day 10, 263 hours, 9.0 mm (Fig. 6). Liver reaching midventral line; yolk largely assimilated. Pectoral fins 0.8 mm long, no fin rays in any fin. Pigmen- tation increasing on dorsal, dorso-visceral, and midventral lines; melanophores on later- al pigment line, opercle, and at pectoral fin base. Postlarval Development Stages 33 and 34: assimilation of yolk, day 12, 311 hours, 10.0 mm. Yolk assimilated, proctodeum open to form anus. Opercles cover two anteriormost gills. Otoliths present in auditory vesicles. Pectoral fins with 3 rays; caudal fin with 3 or 4 rays; no trace of dorsal or anal fins. Spleen forming posterior to liver. Posterior end of notochord (urostyle) up- curved. Pigmentation increased on all aspects except lateral pigment line; melanophores appearing on gas bladder. Stage 35: early postlarva, day 17, 430 hours, 12.0 mm. Caudal fin-rays increased to 7 to 9. Opercles cover gills. Median fin fold thickened and expanded at sites of anal and dorsal fins. Food materials in stomach; feces passing through intestine. All pigmentation excepting lateral pigment hue intensified. Stage 36: dorsal and anal fin-ray rudi- ments, day 40, 960 hours, 15.5 mm (Fig. 7). Fin-ray rudiments in dorsal and anal fins; caudal fin with full complement of ossified rays. Gas bladder constricting into two cham- bers. Melanophores appearing on posterior part of lateral pigment line. Stage 37: middle postlarva, day 48, 1152 hours, 20.0 mm. Rays of dorsal and anal fins partially ossified; pelvic fins rudimentary as gatherings of mesenchyme; caudal fin becom- ing emarginate. Gas bladder two chambered. Urogenital papilla apparent. Mouth terminal; Fig. 6. Late prolarva of razorback sucker, Xyrauchen texanus; legend as in Figures 1 to 5 when applicable, and GB = gas bladder, L = liver, M = mouth, Mp = melanophores, and PF = pectoral fin. December 1982 Minckley, Gustafson: Razorback Sucker Development 559 VPL PFf Fig. 7. Postlarva of razorback sucker, Xijrauchen texanus, at beginning of fin-ray development; legend as in Fig- ures 1 to 6 when applicable, and AFr = anal fin-rays. An = anus, CFr = caudal fin-rays, DFr = dorsal fin-rays DPL = dorsal pigment line, G = gills, LPL = lateral pigment line, N = naris, PFr = pectoral fin-rays, and VPL = ventral pigment line. lips formed, small papillae on both upper and lower lips. Melanophores increasing over body; ossified fin rays and gills acquiring pig- ment; larvae dark dorsally and lighter ven- trally and in eye region. Stage 38: pelvic fin rudiments, day 50, 1200 hours, 20.8 mm. Pelvic fin rudiments in form of small crescentic folds. Stage 39: pelvic fin buds, day 54, 1296 hours, 23.5 mm. Pelvic fin buds in form of thin membranous paddles, not movable. Stage 40: pelvic fin rudiments, day 64, 1536 hours, 27.0 mm. Six pelvic fin-ray rudi- ments within pelvic fin membranes; fin mov- able. Dorsal and anal fin-rays completely ossified. Mouth beginning to shift ventrally, papillae highly developed on lips. Stage 41: ossification of pelvic fin-rays, day 70, 1680 hours, 32 mm. Rays partially os- sified in pelvic fins; median fin membrane persisting ventrally and anterior to pelvic fins greatly reduced; dorsal and anal fins sepa- rated from caudal fin. Stage 42: late postlarva, day 75, 1800 hours, 35 mm (Fig. 8). Pelvic fin-rays ossified. Median fin membrane persisting only be- tween anus and pelvic fins. Full complements of fin-rays in all fins: dorsal, 15-16; caudal, 19; anal, 8; pelvic, 9-9; and pectoral, 13- 15-13-15. Narial flap formed. Alimentary tract looped once to left just posterior to stomach, mouth ventral. Acoustico-lateralis system formed on anterior half of body. Juvenile Development Stage 43: scale rudiments, day 105, 2520 hours, 43.0 mm. Scale rudiments present on ventro-lateral surfaces; median fin membrane eliminated. Stage 44: scales, day 125, 3000 hours, 45.0 mm. Scales distinctly visible except medially on dorsum and ventrum. Stage 45: nuchal keel, day 227, 5448 hours, 90.0 mm (Fig. 9). Scalation completed. Acoustico-lateralis system completed. Nuchal Fig. 8. Late postlarva of razorback sucker, Xijrauchen texanus; legend: LL = lateral line. Lip = papillose lips NF = nasal flap, PgF = pelvic fin, and UgP = urogenital papillus. 560 Great Basin Naturalist Vol. 42, No. 4 Fig. 9. Juvenile of razorback sucker, Xijraiichen texamis, shortly after initiation of nuchal keel. keel evident to touch on predorsum. Except for further development of nuchal keel, evi- dent to the eye at a 250-300 days of age (Fig. 10), morphogenesis is completed. Individuals from the 1974 cohort achieved sexual matu- rity in their sixth year of life (Minckley 1983). Summary Fertilized, water-hardened ova of Xtjraii- chen texanus were 1.8 mm in diameter from females hormone-induced to mature and 2.9 mm from females that matured naturally. Eggs were adhesive for 3 to 4 hours after fertilization. Cleavage was completed in 24 hours at temperatures varying from 13 to 17 C; further development was at 20 C. Gastru- lation occurred at 34 hours. The notocord separated from the mesoderm at 47 hours, and eye rudiments and brain cavities were distinguishable at 49 hours. The tail process formed at 57 hours (2.5 mm). Heart beat be- gan at 83 hours, and blood circulation was es- tabhshed at 117 hours. Embryos began vio- lent flexing and some ruptured tlieir chorions at that time. All hatched between 125 and 131 hours after fertilization. Embryos were 6.8 to 7.3 mm TL at hatch- ing. The yolk sac is tubular, and the head flexes over it at a 45 degree angle to the body axis. The pectoral fin buds, noted at 120 hours (6.8 mm), became paddlelike at 162 hours (8.0 mm), and first were movable at 238 hours (9.0 mm). The continuous, median fin fold first noted on caudal and post- erodorsal areas at 103 hours (5.3 mm), began to develop on the venter (behind the procto- deum) at 142 hours (7.5 mm), and was contin- uous at 162 hours (8.0 mm). The opercles be- gan forming at 162 hours (8.0 mm). The gas bladder first appeared at 238 hours (9.0 mm). The lower jaw became movable and pro- larvae began directed swimming to the sur face to feed at that time; eye pigment com- plete and black. Melanophores developed over mid- and hindbrain and on paired dor- sal, paired dorso-visceral, and unpaired mid- ventral pigment lines between 162 and 238 hours (8.0 and 9.0 mm). There was no lateral Fig. 10. Juvenile of razorback sucker, Xyrauchen texanus, with essentially adult morphology. December 1982 Minckley, Gustafson; Razorback Sucker Dev ELOPMENT 561 pigmentation at 9.0 mm. Melanophores on the dorsum are large and stellate, as also re- corded by Winn and Miller (1952). At 263 hours (9.0 + mm) fin rays were not yet visible in any fin. Yolk was completely assimilated at 311 hours (10.0 mm) and the proctodeum opened to form the anus. The urostyle became up- turned between 263 and 311 hours, and 3 to 4, ventral, caudal fin-rays formed by the last time period. Pectoral fins had developed three rays, but there were no rays in the dor- sal and anal fins. Median fin folds were thick- ened and expanded at the sites of the future dorsal and anal fins at 430 hours (12.0-1- mm), and the opercles fully covered the gills. Dorsal and anal fin-ray rudiments, and a lat- eral pigment line appeared at 960 hours (15.5 mm). The gas bladder had by this time con- stricted toward the two-chambered condi- tion. Pelvic fins appear as swellings of mesen- chyme and the caudal fin becomes emarginate at 1152 hours (20.0 mm). The pelvic fin buds were nonmovable, mem- braneous paddles at 1296 hours (23.5 mm); movement and pelvic fin-rays had appeared at 1536 hours (27.0 mm). Scale rudiments were first noted at 2520 hours (43.0 mm) on ventrolateral body sur- faces. By 3000 hours, lepidogenesis was com- plete on all but the median areas of the dor- sum and ventnmi. The nuchal keel appeared about 5000 hours after fertilization. Acknowledgments This research was supported, in part, by U.S. Fish and Wildlife Service Contract 14- 16-0002-3585 to Arizona State University. We thank personnel at Willow Beach Na- tional Fish Hatchery for assistance in obtain- ing and rearing young of razorback suckers. Permits for collection of wild fish were granted by Arizona Game and Fish Depart- ment. J. P. Collins and R. W. McGaughey read and commented on the manuscript. Literature Cited Balinsky, B. I. 1948. On the development of specific characters in cvprinid fishes. Proc. Zool. Soc. London 118:335-344. Douglas, P. A. 1952. Notes on the spawning of the razorback sucker, Xyraiichen texanus (Abbott). Cahfornia Fish and Game 38(2): 149-155. FuiMAN, L. A., AND D. C. WiTMAN. 1979. Descriptions of catostomid fish larvae: Catostomus catostomus and Moxostoma enjthruntm. Trans. Amer. Fish Soc. 108(6):604-619. FuiMAN, L. A., AND J. R. Trojnar. 1980. Factors affect- ing egg diameter of white sucker {Catostomus commersoni). Copeia 1980(4):699-704. Long, W. L., and W. W. Ballard. 1976. Normal em- bryonic stages of the white sucker, Catostomus commersoni. Copeia 1976(2):342-351. Minckley, W. L. 1983. Status of the razorback sucker, Xijrauchen texanus, in the lower Colorado River basin. Southwest. Nat. 28(2): In press. PisTER, E. P. 1981. The conservation of desert fishes. Pages 411-446 in R. J. Naiman and D. L. Soltz, eds.. Fishes in North American deserts. John Wiley and Sons, New York. Ryder, J. A. 1885. On the development of viviparous osseous fishes. Proc. U.S. Nat. Mus. 85:128-162. Stewart, N. H. 1926. Development, growth and food habit of the white sucker, Catostomus com- mersonii LeSueur. Bull. U.S. Bur. Fish. 42:147-183. Ta VOLGA, W. N. 1949. Embryonic development of the platyfish (Platypoecilus), the swordtail (Xipho- phorus), and their hybrids. Bull. Amer. Mus. Nat. Hist. 94(4): 161-230. ToNEY, D. P. 1974. Observations on the propagation and rearing of two endangered fish species in a hatch- ery environment. Proc. Ann. Conf. West. Assn. State Game Fish Comm. 54:252-259. Winn, H. E., and R. R. Miller. 1954. Native postlarval fishes of the lower Colorado River basin, with a key to their identification. California Fish and Game 40(3):273-285. BEHAVIOR AND HABITAT PREFERENCES OF RING-NECKED PHEASANTS DURING LATE WINTER IN CENTRAL UTAH Jeffrey G. Skousen' and Jack D. Brotherson' Abstract.- Ring-necked pheasant behavior and habitat preferences were studied during late February along benches of the Wasatch Mountains in central Utah. Seven behavioral categories were used to classify pheasant activ- ities during three time periods of the day. Eating, alert, and movement behavior were the most frequent activities during all periods of the day. Significant differences (p<.01) were found between morning and midday behavior and between midday and evening behavior. Pheasants fed in open areas during morning hours then retreated into heavy cover during midday periods. The birds remained in heavy cover until late afternoon. Pheasants then moved away from heavy cover into semiopen areas to feed as evening approached. The ring-necked pheasant {Phasianus col- chicus) is a native of eastern Asia. Its first successful importation into the United States was in Oregon in 1881 (Allen 1956). Later, in 1888, the birds were also successfully trans- planted into the eastern part of the United States (Rue 1980). Pheasants were introduced into the state of Utah about 1890 (Rawley and Bailey 1972). Distribution of pheasants in Utah has increased so that nearly all suitable habitat is occupied. This habitat is primarily within the irrigated farmlands of the state, which comprises about 2 to 4 percent of the total land area (Olsen 1977). The pheasant population of Utah reached a peak in the 1950s and has steadily decreased. This down- ward trend has been a result of habitat dete- rioration (Nish 1973). Pheasant distribution is primarily affected by the habitat, soil, and climate of an area. Christensen (1951) found that together soils and climate determined whether or not pheasants occurred in an area of Missouri. Dale (1956) has suggested that a combination of high temperature and high humidity have probably inhibited the spread of pheasants into the southeastern United States. Soils and climate restrict pheasant distribution because climate controls soil development and food quality. Climate is especially important dur- ing the nesting season because hens leave their eggs for a period of time after laying, thus exposing them to climatic conditions (Graham and Hesterberg 1948). Pheasant eggs generally show reduced hatchability as temperatures increase (Yeatter 1950). Wood and Brotherson (1982) found that nesting sites were specifically dependent upon total hving ground cover and high amounts of cover sur- rounding the nesting cavity. Soils seem to influence pheasant distribu- tion through their effects on vegetation. In the United States much of the fertile soil is cultivated with corn and other grains, which have been shown to be preferred pheasant food (Leedy and Hicks 1945). Christensen (1951) reported that the- distribution of pheasants in Missouri coordinated with high- ly fertile soils used for agriculture. Edminster (1954) agreed that pheasants preferred agri- cultural land that produced grain. Pheasants showed similar distribution patterns in Wis- consin (Hine 1964), Ohio (Leedy and Hicks 1945), and Illinois (Kimball et al. 1956). In Montana and Utah, conversion of grain-pro- ducing acreage to hayfields has resulted in a decline of pheasant densities (Weigand 1973, Nish 1973). Winter cover and nesting habitat are the two major deficiencies in pheasant habitat in the Great Lake States (MacMullan 1961, McCabe et al. 1956), the Dakotas (Kimball et al. 1956), the northwest (Lauck- hart and McKean 1956), New York (Peny 1946), and California (Hart et al. 1956). In areas of Utah where adequate cover is lacking, winter kill of pheasants has been shown to be high (Yeager et al. 1956). Wood and Brotherson (1981) found that in most areas of Utah over- 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 562 December 1982 Skousen, Brotherson: Pheasant Behavior 563 winter pheasant mortality was not due to winter weather. Pheasants are omnivorous; eating waste grains, weed seeds, green vegetation, and nu- merous insects (Rue 1980, Olsen 1977, Raw- ley and Bailey 1972). Different proportions of these foods are eaten depending on area, location (Cottam 1929, Swenk 1930, Gigstead 1937, Fried 1940, Trautman 1952, Korschgen 1964), and the age of the bird (Rue 1980, Ed- minster 1954). In Utah, adult pheasant diets are comprised mostly of grain (36.7 percent) and vegetation (20.4 percent) (Cottam 1929). In the midwest, 70 to 85 percent of the total diet of adult pheasants is cultivated grains (Dalke 1937, Trautman 1952, Korschgen 1964, Fried 1940, Gigstead 1937, Swenk 1930). Juvenile birds, for the first three weeks, eat insects (Rue 1980, Edminster 1954, Dalke 1937, Loughrey and Stinson 1955) then switch to vegetation after that. Few studies have been done on actual daily behavioral patterns of the ring-necked pheas- ant. Burger (1966) studied aggressive territo- rial behavior of male ring-necked pheasants during the mating season. Kuck et al. (1970) reported on the renesting behavior of hen pheasants in South Dakota. This paper is con- cerned with habitat preferences and daily be- havior patterns of pheasants in central Utah during late February. Study Area The study area (40 ha) is located on a bench of the Wasatch Mountains near Provo, Utah. Thirty percent of the area is a shrub study plot used by the Intermountain Forest and Range Experiment Station, 20 percent is Table 1. Activities associated with each behavior category. Category Behavior Eating Resting Grooming Aggressive Alert Movement Comfort Head is down, pecking material on the ground. Eyes closed and relaxed. Fluffing feathers. Cause another bird to move or pecking other birds. Watching, alert, and ready to fly. Looking at something moving or making noises close by. Walking, running, or flying. Looking without being alert and not at anything in particular. Scratching. a mature apricot orchard (used for pasture) and 50 percent is native rangeland. The area has a western aspect with loamy cumulic cal- cixeroll soils. Slopes are generally 3 to 10 percent and elevation is approximately 1400 m (4600 ft). A housing tract is located on the lower edge of the study area, and one paved and heavily used road runs through the cen- ter of the area. Methods Five days of 4 to 8 hours each were spent observing pheasants during 20-24 February 1982. Days during which the study took place were generally clear and sunny with some scattered clouds. Approximately 15 to 25 cm (6-10 in) of snow was on the ground when the study was initiated but melted be- fore completion. Temperatures on these days ranged from 35 F in the morning hours to 60 F during afternoons. Observations were made with the aid of binoculars and a 20 power spotting scope. One individual was chosen (a male normally) and observed closely. At one- minute intervals, the individual was classified as to his behavior at that moment. The be- havior classes considered were: eating, rest- ing, grooming, aggressive, alert, movement, and comfort behavior (Table 1). In between the minute evaluations, interaction between the selected individual and others, as well as other activities that influenced the group, were recorded. Habitat utilized by the birds was also noted and later classified into habi- tat types. Results and Discussion Pheasant behavior was variable during the day but was very similar during morning and Table 2. Behavioral classes and percent of time spent in each class during three time periods. Morning Midday Evening Behavioral 6:00-10:00 10:00-4:00 4:00-8:00 class (percent) (percent) (percent) Eating 47.5 7.0 51.5 Resting 0.0 1.0 0.0 Grooming 3.5 4.0 0.5 Aggressive 2.5 2.0 0.5 Alert 3L5 50.0 22.0 Movement 14.5 32.0 20.5 Comfort 0.5 4.0 6.5 564 Great Basin Naturalist Vol. 42, No. 4 evening hours (Table 2). Approximately 50 percent of the time spent during these peri- ods was in eating, and another 40 to 45 per- cent of the time was divided between alert (25 percent) and/ or movement (20 percent) behavior (Table 2). Grooming, aggressive, and comfort behavior contributed the bal- ance of the time spent (6-7 percent). How- ever, during midday periods, most of the time was spent in being alert (50 percent) and/ or moving (32 percent) (Table 2). Eating behavior during this period was reduced to 7 percent. Evening behavior of pheasants, though similar to morning periods, seemed to be less affected by noises made by man. There was a highly significant difference (p<.01) in the amount of time spent in each behavior category across a day. Chi-square tests showed no significant difference in be- havior patterns between morning and eve- ning periods of the day. However, significant differences (p<.01) were observed between morning and midday behavior and between midday and evening behavior (p<.01). Eat- ing, alert, and movement behavior were the most frequent activities during all times of the day. Spearman's rank correlation (Snede- cor and Cochran 1968) showed no significant differences between rankings of observed be- havior during morning, midday, and evening. Habitats utilized by the pheasants were classified into 5 types. Table 3 shows these habitat types and their major associated plant species. Habitat preference varied between periods of the day (Table 4). During morning hours, pheasants were observed to be in areas that offered little cover and where pasture grasses were available (Tables 3, 4). Numer- ous insects were observed in the grass pas- ture. The birds may have been eating the in- sects rather than the vegetation. They moved to areas of heavier cover (ditch banks and fence rows) as activities of man increased and as disturbances from other animals increased. During midday periods, all pheasants ob- served were found in fence rows or ditch Table 3. Species list in each habitat type. Orchard grass Orchard grass Forest apricot apricot pasture Wooded Service Species pasture fence row fence row Ditch banks plots Agoseris glauca X X X Agropyron repens X X Agropyron spicatum X X X X X Ambrosia psilostachya X X X X X Artemisia tridentata X Bromus tectorum X X X X X Ailanihus altissima X Chrysothamniis nauseosus X Cirsiiim arvense X X Cirsiiim vulgare X X Desciirainia sophia X X X X Dactylis glomerata X X X X Erodium cicutarium X X X Grindelia squarrosa X X Kochia prostrata X Kochia scoparia X X X X Linaria vulgaris X Penstemon spp. X Petradoria pumiki X X Poa secunda X X X X X Primus armenica X X Populus angiistifolia X Populus fremontii X X Rosa woodsii X > Robinia pseudoacacia X Salix fragilis X Salsola kali X Sy ringa vulgaris X Ulmus pumila X Xanthium spp. X X December 1982 Skousen, Brotherson: Pheasant Behavior 565 Table 4. Habitat and cover preferences and percent of time spent in each habitat. Habitat type Morning (percent) Midday (percent) Evening (percent) Orchard grass apricot pasture 29 17 0 Orchard grass apricot pasture fence row 62 0 0 Wooded fence rows 0 21 0 Ditch banks 9 62 54 Forest Service plots 0 0 46 banks (Table 4). Evening hours were spent along ditch banks and shrub plots that of- fered some cover but which were generally more open than the areas frequented at midday. Movement from open areas in the morning to areas of heavier cover (i.e., ditch banks and fence rows) depended on the amount and kind of disturbances in the immediate area. Cars starting and children walking to and from school generally caused the pheasants to move slowly into heavier cover. Pheasants ran and flew as crows passed overhead. Mid- days were spent in heavy cover and the birds did not move unless flushed. Evenings were generally spent feeding and moving away from heavy cover while being very cautious and aware of the activities around them. Literature Cited Allen, D. L., ed. 1956. Pheasants of North America. Stackpole Co., Harrisburg, Pennsylvania, and Wildlife Management Institute, Washington D.C. Burger, G. V. 1966. Observations on aggressive behavior of male ring-necked pheasants in Wisconsin. J. Wildl. Manage. 30(l):57-64. CoTTAM, C. 1929. The status of the ring-necked pheas- ant in Utah. Condor 31(3):117-123. Christensen, D. M. 1951. History and status of the ring- necked pheasant in Missouri. Missouri Cons. Commission. Dale, F. H. 1956. Pheasants and pheasant populations. Pages 1-42 in Allen 1956. Dalke, p. L. 1937. Food habits of adult pheasants. Pages 139-142 in McAtee 1945. Edminster, F. C. 1954. American game birds of fields and forest. Charles Scribner's Sons, New York. Fried, L. A. 1940. The food habits of the ring-necked pheasant in Minnesota. J. Wildl. Manage 4(l):27-.36. GiGSTEAD, G. 1937. Habits of Wisconsin pheasants. Wil- son Bull. 49(l):28-34. Graham, S. A., and G. Hesterberg. 1948. The influence of climate on the ring-necked pheasant. J. Wildl. Manage. 12(1):9-14. Hart, C. M., B. Glading, and H. T. Harper. 1956. The pheasant in California. Pages 90-158 in Allen 1956. HiNE, R. L. 1964. Status of farm game in Wisconsin. A Report of the Farm Game Working Group of the Natural Resources Committee of State Agencies Land Subcommittee. Madison, Wisconsin. Kimball, J. W., E. L. Kozicky, and B. A. Nelson. 1956. Pheasants of the plains and prairies. Pages 204-263 in Allen 1956. Korschgen, L. J. 1964. Foods and nutrition of Mi.ssouri and Midwestern pheasants. Trans. N. Am. Wildl. Conf. 29:159-181. KucK, T. L., R. B. Dahlgren, and D. R. Progulske. 1970. Movements and behavior of hen pheasants during the nesting season. J. Wildl. Manage. 34(3);626-630. Lauckhart, J. B., AND J. W. McKean. 1956. Chinese pheasants in the Northwest. Pages 4.3-89 in Allen 1956. Leedy, D. L., and L. E. Hicks. 1945. The pheasants of Ohio. Pages 57-130 in McAtee 1945. Loughrey, a. G., and R. H. Stinson. 19.55. Feeding habits of juvenile ring-necked pheasants on Pelee Island, Ontario. Canadian Field Nat. 69(2):59-65. MacMullan, R. a. 1961. Ring-necked pheasant habitat management in the United States. Trans. N. Am. Wildl. and Nat. Resour. Conf. 26:268-272. McCabe, R. a., R. a. MacMullan, and E. H. Dustman. 1956. Ringneck pheasants in the Great Lakes region. Pages 264-356 in Allen 1956. NisH, D. H. 1973. Pheasant habitat trends in Utah-can they be reversed? Proc. Pheasant Habitat Sym- posium, Salt Lake City, Utah. 11 July 1973. Olsen, D. W. 1977. A literature review of pheasant hab- itat requirements and improvement methods. Utah Div. of Wildl. Res. Pub. No. 77-7. 144 pp. Perry, R. F. 1946. An appraisal of pheasant abundance in New York State during 1945 and some of the factors responsible for the recent decline. Trans. N. Am. Wildl. Conf. 11:141-155. Rawley, E. v., and W. J. Bailey. 1972. Utah upland game birds. Utah Div. of Wildl. Res. Pub. No. 63- 12. 31 pp. Rue, L. L. 1980. Wildlife profiles: ringneck pheasant. American Hunter Vol. 8, No. .3, March 1980, pp. 40-43. Snedecor, G. W., and W. G. Cochran. 1968. Statistical Methods. 6th ed. Iowa State Univ. Press, Ames, Iowa. 593 pp. Swenk, M. H. 19.30. The food habits of the ring-necked pheasant in central Nebraska. Nebraska Agric. Expt. Sta., Research Bull. 50. ,33 pp. Trautman, C. G. 1952. Pheasant food habits in South Dakota. Tech. Bull. No. 1, South Dakota Dept. Game, Fish and Parks. Weigand, J. P. 1973. Pheasant population and habitat trends in Montana. Proc. Pheasant Habitat Sym- posium, Salt Lake City, Utah. 11 July 1973. 566 Great Basin Naturalist Vol. 42, No. 4 Wood, A. K., and J. D. Brotherson. 1981. Overwinter Yeager, L. E., J. B. Low, and H. J. Figge. 1956. Pheas- survival of ring-necked pheasants in Utah. Great ants in the arid Southwest. Pages 159-203 in Al- Basin Nat. 41(2):247-250. len 1956. 1982. Microhabitat and nest site selection by Yeatter, R. E. 1950. Effects of different preincubation ring-necked pheasants. Great Basin Nat. temperatures on the hatchability of pheasant 41:457-460. eggs. Science 112(2914):529-530. RHYTHM OF FECAL PRODUCTION AND PROTEIN CONTENT FOR BLACK-TAILED JACKRABBITS William D. Steigers, Jr.', Jerran T. Flinders', and Susan M. White''- Abstract.- The cyclical phenomenon of soft and hard formation of feces in desert black-tailed jackrabbits (Lenus califormcus deserticola, Mearns) was investigated. Sixty-nine blacktails were shot between 0525 hours and 1508 hours over a 4-day period. The average age was 314 days for all black-tails. The large intestine was removed and linear measurements taken. Overall length of the large intestine averaged 163.3 cm; mean length of the colon was 51 5 cm arid average length of the rectum was 111.8 cm. Moisture content for soft and hard pellets averaged 79 5 percent and 74.1 percent, respectively. Protein content of soft and hard pellets averaged 45.8 percent and 14 3 percent respec- tively^ The black-tails began the transition from hard to soft pellets as early as 0414 hours, and nearly all had cnjm- pleted the reverse transition by 1508 hours. Female black-tails began both' early morning and afternoon transitions significantly later than the males. Reasons for the apparent sexual partitioning are proposed. It has long been known that the domestic rabbit produces both hard and soft fecal pel- lets (Morot 1882). The hard pellets are the normal waste product of the digestive tract and are the type of feces found in the field. The soft "pellets" are a special product of the cecum, and are reingested directly from the anus and swallowed whole. The position assumed by the black-tailed jack rabbit when reingesting soft pellets is described by Lech- leitner (1957). The method whereby a rabbit can pass two kinds of feces, soft and hard pellets, has been conjectured by Eden (1940) and Thacker and Brandt (1955). The latter suggest hard fecal pellets are formed by material that has passed through the base of the cecum with- out being mixed with the main content of the cecum. The soft feces are formed by empty- ing the major portion of the cecum, in a cy- clic manner, by a strong contraction of the spiral muscle (Thacker and Brandt 1955). The composition of the soft feces is comparable to that of the cecal contents in protein, crude fi- ber, and other proximate nutrients (Eden 1940, Olsen and Madsen 1944). Cececto- mized rabbits do not excrete typical soft feces (Hemdon and Hove 1955). Although the hard pellets exit the body as firm, nearly spherical excreta, the soft feces are generally excreted as clusters of soft, moist, pellets with a distinctive sheen. Soft feces usually are higher in protein than hard feces (Herndon and Hove 1955). Soft pellets consist largely of bacteria surrounded by a proteinaceous membrane deposited posterior to the colon (Griffiths and Davies 1963). Several authors have reported the cycle or rhythm of production of hard and soft pellets in rabbits. Although Southern (1942) reported the frequency of coprophagy (or cecaphagy) in the wild rabbit as twice daily, most au- thors suggest a single such daily period (Meyer 1955, Watson and Taylor 1955, Lech- leitner 1957). Spencer (1955) studied rhythms of reingestion in white-tailed jackrabbits {Lepiis townsendii), snowshoe hares (L. amer- icanus), and New England cottontails {Syl- vilagus transitionalis). Hansen and Flinders (1969) suggested that reingestion in white- tailed jackrabbits, black-tailed jackrabbits, snowshoe hares, and European hares takes place in the late morning hours. Thacker and Brandt (1955) found that the excretion of hard and soft feces by domestic Dutch rabbits was a consistent daily phenomenon both as to time and quantity. Lechleitner (1957) re- ported the percent of soft feces in the recta of 160 black-tailed jackrabbits killed period- ically throughout every month of the year. Although the exact time of death was not known, it had been approximated for the 160 'Wildlife and Range Resources Graduate Program, 401 WIDE, Brigham Young University, Provo Utah 84602 'Present address: Native Plants, 400 Wakara Way, Salt Lake City, Utah 84108. 567 568 Great Basin Naturalist Vol. 42, No. 4 black-tails. However, no known study re- ported to date has attempted to quantify the exact periodicity of soft and hard feces for- mation in wild populations of black-tailed jackrabbits. The purpose of this study was to quantify the exact time periods of production as well as transition periods for hard and soft feces in wild populations of black-tailed jackrabbits in north central Utah. Study Area This study was conducted in two areas in north central Utah. Rabbits were sampled from rangelands near Fountain Green, San- pete County, Utah, and from the U.S. Forest Service Benmore Experimental Area near Vernon, Tooele County, Utah. The elevation of the study areas are approximately 1830 m and 1770 m, respectively. Each receives about 35 cm of average annual precipitation, with most of this in the form of snow from October through May. The woody vegetation of both areas is dominated by rubber rabbitbrush {Chrijso- thamnus nauseosus) and big sagebrush {Arte- misia tridentata). Prominent grasses include bluebunch wheatgrass {Agropyron spicatum), fairway wheatgrass (A. cristattim), and Sand- berg bluegrass {Poa sandbergii). Methods Desert black-tailed jackrabbits were col- lected with shotguns on four days during March 1980. Predawn collections were con- ducted with the aid of a spotlight. The first 59 black-tails were collected from the Foun- tain Green study area: 8 black-tails on 8 March, 19 blacktails on 15 March, and 32 blacktails on 18 March. An additional 10 black-tails were taken 22 March from the Benmore study area. All black-tails were taken opportunisti- cally, and the exact time of death to the near- est minute was recorded for each hare shot. An attempt was made to sample black-tails during all time periods between initiation of the hard-to-soft pellet transition in early morning to the soft-to-hard pellet transition in the early afternoon. Entrails of the black- tails were removed within 3 hours after being shot, and frozen. The left eyeball was removed for age de- termination using the eye lens-weight meth- od and age curve described by Tiemeier (1965). The eyeball was held in 10 percent buffered formalin for one week. The lens was then removed from the eyeball and placed in an oven at 90 C. The lens was considered dry after 48 or more hours, when repeated weighings to 0.001 g resulted in no additional weight loss. In the laboratory, the portion of the in- testinal tract from the anus to the juncture of the small intestine and ileocecal valve were separated by severing the mesentery. The overall length of the large intestine from the ileocecal valve to the anus was measured, as well as the length from the ileocecal valve to the taenia coli muscle. Fecal samples of both hard and soft pellets were taken, if present, from the rectum (between taenia coli muscle and anus). Hard pellets could not be accu- rately separated from soft pellets in the large intestine until the feces passed the taenia coli muscle. If a transitory condition between hard and soft pellets was noted, the length of the rectvim between the transition and taenia coli muscle was measured. The precise time the transition occurred at the taenia coli muscle was calculated by backdating the transition for a 45 cm per hour rate of pas- sage. Fecal samples were immediately weighed to 0.001 g, and then placed in a drying oven at 83 C for 24 hours to deter- mine moisture content. Within approximately one month, the fe- cal samples were again oven dried, cooled in a dessicator, ground, and analyzed for nitro- gen content using the macro-Kjeldahl tech- nique (Black et al. 1965). Samples of feces <1 g were pooled with fecal samples from black-tails of the same sex that were collect- ed at approximately the same time. Results and Discussion Measurements The 69 (33 males, 36 females) black-tailed jackrabbits were collected between 0525 hours and 1508 hours. The average age for all black-tails was 314 ± 141 days (SD) {N = 67, range 153-746 days). The average age for males was 315 ±142 days (iV=31, range December 1982 Steigers et al.: Black-tailed Jackrabbit 569 n u 150 250 350 450 550 650 750 200 300 400 500 600 700 AGE (DAYS) Fig. 1. Histogram illustrating age distribution in days of black-tailed jackrabbits collected for analysis. 153-718 days) and for females was 313 ± 143 days {N=36, range 169-746 days). The aver- age age by 50-day intervals for black-tails in our sample is illustrated in Figure 1. Black- tails bom during 1979 comprised 84 percent of the sample; all remaining black-tails were bom during 1978. Seventy-two percent of all black-tails collected were between 175 and 325 days of age and were born during the pe- riod June through September. A breakdown by year and month for black-tails born in 1978 and 1979 is represented in Figure 2. The average overall length of the large in- testine was 163.2 ±17.2 cm (iV=66, range 134-212 cm). The mean length of the colon was 51.5 ±5.0 cm (iV=66, range 43-72 cm) and of the rectum was 11 1.7 ±14.6 cm (iV = 66, range 85-143 cm). Correlation be- tween the measurements of overall length of the large intestine (r = 0.31, P < 0.05), colon length (r = 0.25, P < 0.05), and rectal length (r = 0.29, P < 0.05) with age of the black-tail was relatively low. However, correlation be- tween the overall length of the large intestine and colon length (r = 0.62, P < 0.001) and the overall length of the large intestine and rectum length (r = 0.96, P < 0.001) was much higher. The correlation between colon length and rectal length was r = 0.38 {P < 0.005). Average percent moisture of soft pellets from 58 samples collected from the rectum was 79.5 ±1.8 percent (range 74.2-83.0 per- cent). Average percent moisture of hard pel- lets from 20 samples collected from the rec- 15l ia S5. n ^^'^feb'^'^"apr^'^^jun>J^'-augS^^ct'^0Vdec month of birth Fig. 2. Distribution of months of birth for black-tailed jackrabbits collected for analysis that were born during 1978 and 1979. tum was 74.1 ±2.8 percent (range 67.8-78.8 percent). The average moisture content for soft pellets was significantly greater (P < 0.001) than that of hard pellets using an un- paired T-test. This comparison included 12 samples where hard and soft pellets were re- moved from the same rectum. This indicates a significant change in moisture content asso- ciated with the rapid change in the type of pellet produced at the taenia coli muscle. Protein Protein content (percent N X 6.25) of soft and hard pellets averaged 45.8 ±5.9 percent (N = 8, range 35.0-51.5 percent) and 14.3 ±5.6 percent {N=9, range 7.1-23.3 per- cent), respectively. Derived protein values compare favorably with those reported by other authors. Herndon and Hove (1955) re- ported 41.9 percent protein for soft pellets and 14.8 percent protein for hard pellets in experimental California-white rabbits {Oryctolagus cuniculus). Griffiths and Da vies (1963) estimated soft feces of rabbits con- tained 24.4 percent protein, 81 percent of which was in the form of bacterial cells. They reported bacterial cells composed 56 percent of the dry weight of soft feces. Other comparative estimates of protein content in the soft feces of domestic rabbits were 37.8 percent (Huang et al. 1954) and 28.5 percent 570 Great Basin Naturalist Vol. 42, No. 4 Males (N = 11) Females (N = 14) lOOr 90 80 70 60 50 40 30 m 33 o u m ■ I • \ -0 o O t c o ■D I- m 20 ■ 10 : 4 i I 1 I • I 10 12 14 16 18 TIME OF DAY (HOURS) Fig. 3. Percent of male and female black-tailed jack- rabbits producing soft pellets at the taenia coli muscle in the large intestine. for soft feces and 9.2 percent for hard feces (Kulwich et al. 1953). Thacker and Brandt (1955) reported 37.4 ±3.2 percent protein for soft feces and 18.7 ±1.7 percent protein for hard feces in the Dutch rabbit. Pellet Transition Periods When a transition zone was found in the rectum of a black-tail, the degree of dis- tinctness between the two types of pellets was noted. In our samples, 17 of 19 transition zones were very well defined and distinct, of- ten with immediately adjacent pellets of the two types. In the other two cases, the transi- tion zone included up to 10 cm of semihard pellets separating the hard and soft pellet types. Distinct transition zones were found for both early morning and afternoon periods of pellet transition. Male black-tailed jackrabbits had initiated the transition from hard to soft pellets as early as 0414 hours (Fig. 3). The transition period for both sexes ranged from 0414 to ap- proximate completion by 0715 hours. Males sampled were complete in the initiation of their transition by 0500 hours, whereas fe- males sampled were not all complete by 0700 hours (Fig. 3). All black-tails had made the transition to the production of soft pellets by 0730 hours. Soft pellets made up the entire composition of the rectum in all black-tails by 0915 hours. Several male black-tails began the transi- tion from soft to hard pellets as early as 1201 hours, or slightly before (Fig. 3). Female black-tails tended to initiate the same transi- tion later in the afternoon. The earliest noted transition for a female was 1335 hours and most females initiated the transition between 1400 to 1430 hours. Most males had com- pleted the transition to hard pellets by 1330 hours, but one male had started as late as 1450 hours (Fig. 3). Two females had not in- itiated the early afternoon transition by 1500 hours. Feces in the rectum of two males were composed entirely of hard pellets as early as 1408 hours and 1414 hours, respectively. No females had fully replaced the soft pellets in the rectum with hard pellets by 1500 hours. The findings presented here generally sup- port those reported by other authors (Spen- cer 1955, Watson and Taylor 1955, Lechleit- 1957, Hansen and Flinders 1969). ner Spencer (1955) found half the animals killed from 0900 hours to 1700 hours had soft pel- lets in their stomachs. There was evidence of soft pellets in stomachs of 21 of 25 animals killed at 1300 hours (Spencer 1955). Soft pel- lets were produced in all black-tailed jack- rabbits collected from 0800 hours to 1000 hours in California, with evidence of the hares starting to reingest them (Lechleitner 1957). Amorphous pellets of the European hare {Lepiis europaeus) were found in the recta of hares killed between 0600 hours and 1600 hours; all animals killed between 0800 hours and 1200 hours contained soft pellets in their recta (Watson and Taylor 1955). Lech- leitner (1957) reported soft feces being pro- duced and swallowed during the daylight hours, starting at 0600 hours and being re- placed by hard feces by 2000 hours. How- ever, the exact time of death was not known for the black-tails used in the sample, and December 1982 Steigers et al.: Black-tailed Jackrabbit 571 sample sizes were minimal during the early morning hours. With the knowledge of the exact time of death, the periods of transition in pellet types in the black-tailed jackrabbit were more precise in our study. Lechleitner (1957), in reporting the cycle of reingestion and production of soft pellets in the black- tailed jackrabbit in California, found the pe- riod of occurrence of soft pellets in the rec- tum may extend to as late as 2000 hours. Our findings do not support his conclusion. We suggest that, aldiough pellets may be found in the rectum as late as 1600 hours or 1700 hom-s, the transition to hard pellets in the af- ternoon is essentially completed by 1330 hours for males and 1430 hours for female black-tails. Our data support the theory that males begin the transition from hard to soft pellets earlier in the morning than do fe- males. This relationship was significant {P < 0.05) using the Mann- Whitney- Wilcoxon 2- independent sample procedure. Females in- itiated the transition from soft to hard feces significantly later (P < 0.01) in the afternoon than males. Lechleitner (1957) found no evi- dence that indicated any seasonal trend of reingestion in the black-tailed jackrabbit. Thus, this relationship may in fact be a year- long phenomenon. The stimulus-response mechanisms causing the apparent dimorphism in behavioral phys- iology between the sexes is not known. How- ever, reasons why this occurs may include partitioning of temporal feeding activities to reduce intersexual competition. This may be especially critical near the end of the winter season when forage above the snow level may be limiting, and when females are in late gestation or lactating. Intraspecific com- petition is maximal under peaking population cycles as well as during climatic seasons of stress. Intersexual segregation of temporal feeding cycles may tend to reduce this com- petition. Because of the higher energy re- quirements of gestation and lactation, female black-tails may feed later into the morning hours than males. Lactating females may suckle leverets in the early morning hours, and this demanding activity may help delay the production of soft pellets. In addition, fe- males may reingest soft pellets later into the afternoon than males, thereby maximizing feeding efficiency and digestability of in- gested forage. Acknowledgments We sincerely thank the following for their time, effort, and private funding so gra- ciously donated to the collection of jackrab- bits for this study: D. A. Barnum, J. S. Cas- trale, F. V. Luke, S. I. Steigers, and especially C. L. Elliott. We also thank H. Cheung for manuscript preparation and J. S. Castrale for his critical review of this manuscript. Figures were drawn by S. I. Steigers and N. P. Hebbert. Literature Cited Black, C. A., D. C. Evans, J. L. White, and others. 1965. Methods of soils analysis. Part I, Agronomy No. 9., Am. Soc. Agron. and Am. Soc. Testing Mater., Madison, Wisconsin. 802 pp. Eden, A. 1940. Origin of night feces. Nature 145:628-629. Griffiths, M., and D. Davies. 1963. The role of soft pellets in the production of lactic acid in the rab- bit stomach. J. Nutr. 80:171-180. Hansen, R. M., and J. T. Flinders. 1969. Food habits of North American hares. Sci. Ser. No. 1, Colorado State Univ., Fort Collins. 18 pp. Herndon, J. E., AND E. L. Hove. 1955. Surgical removal of the cecum and its effect on digestion and growth in rabbits. J. Nutr. 57:261-270. Huang, T. C, H. E. Ulrich, and C. M. McCay. 1954. Antibiotics, growth, food utilization, and the use of chromic oxide in studies on rabbits. J. Nutr 54:621-630. KuLwicH, R., L. Struglia, and p. B. Pearson. 1953. The effect of coprophagy on the excretion of B vitamins by the rabbit. J. Nutr. 49:639-645. Lechleitner, R. R. 1957. Reingestion in the black-tailed jack rabbit. J. Mammal. 38:481-485. Meyer, K. 1955. Coprophagy in the European rabbit (Oryctolagus cuniculus) in Australia. Austr. J. Zool. 3:336-345. Morot, C. 1882. Des pelotes stomacales des leporides. Mem. Soc. Centr. Med. Vet. 12:137. In J. S. Wat- son. 1954. Reingestion in the wild rabbit, Orycto- lagus cuniculus (L.). Proc. Zool. Soc. Lond. 124:615-624. Olsen, H. M., and H. Madsen. 1944. Investigations on pseudorumination in the rabbit. Vidensk. Medd. fra Dansk natrh. Foren. 107:37. In E. J. Thacker and C. S. Brandt. 1955. Coprophagy in the rab- bit. J. Nutr. 55:375-385. Southern, H. N. 1942. Periodicity of refection in the wild rabbit. Nature 149:553-554. Spencer, J. L. 1955. Reingestion in three American spe- cies of Lagomorphs. Lloydia. 18:197-199. Thacker, E. J., and C. S. Brandt. 1955. Coprophagy in the rabbit. J. Nutr. 55:375-385. Tiemeier, O. W. 1965. Bionomics. Pages 5-40 in C. P. Wilson, director. The black-tailed jack rabbit in Kansas. Kansas State Univ., Ag. Exp. Sta. Tech Bull. 140. 75 pp. Watson, J. S., and R. H. Taylor. 1955. Reingestion in the hare Lepus europaeus. Pol. Sci. 121:314. PRAIRIE DOG COLONY ATTRIBUTES AND ASSOCIATED VERTEBRATE SPECIES Tim W. Clark', Thomas M. Campbell IIP, David G. Socha', and Denise E. Casey' Abstract.- A survey of colony attributes and associated vertebrates on black-tail {Cijnomijs hidovicianus), Gun- nison's (C. gimnisoni), and white-tail (C. leucurus) prairie dogs was made. A belt transect 1.6 km wide and 13,.3.34 km long from Hobbs, New Mexico, to the Utah-Wyoming state line was surveyed. There were 47 colonies located (4760 ha comprising 2.2 percent) in the belt. Intercolony distances varied significantly. Three black-tail towns averaged 33 ha in area (SD = 26, range 10-61), II Gunnison's averaged 46 ha (SD = 43, range 16-150), and 33 white-tail towns av- eraged 125 ha (SD=200, range 0.2-958). Badger activity was positively and significantly correlated to colony size and number of burrow openings on Gunnison's and white-tail towns. There were 107 vertebrate species and sub- species (one amphibian, 25 reptiles, 51 birds, 30 mammals) observed on prairie dog colonies. Results of our surveys are compared with prairie dog studies elsewhere. The role of prairie dogs and relationships to some vertebrates spe- cies are discussed. This paper describes results of a survey of colony characteristics and associated verte- brate species for three prairie dog species in Utah, Colorado, and New Mexico. Study Area Prairie dog colonies surveyed were in a belt transect (1.6 km wide and 1334 km long) beginning near Hobbs, New Mexico, and ending on the Utah-Wyoming state line (Fig. 1). The transect generally followed an exist- ing pipeline corridor. The prairie dog species encountered were: black-tail {Cynomys lu- dovicianus), Gunnison (C. gunnisoni), and white-tail (C. leucurus). These species collec- tively occupied many vegetation-phys- iographic types. Overall, black-tail colonies were in shortgrass prairie with Bouteloua sp. and Buchloe dactyloides with scattered Opuntia imhricata. Gunnison colonies were associated with Juniperus monosperma, shrubs, and O. imbricata, as well as a variety of forbs and grasses. White-tail colonies had an overstory of Artemisia sp. and a diverse understory of forbs and grasses. Methods All prairie dog colonies were aerially lo- cated and mapped onto U.S. Geological Sur- vey 7.5 and 15 minute maps. Beginning in June 1980, near Hobbs, New Mexico, and working north to Wyoming, each town was visited, precisely mapped, and inventoried in detail. Surveys followed guidelines designed for black-footed ferret {Mustela nigripes) searches (Clark and Campbell, in prepara- tion), and allowed for a concomitant general survey of vertebrate species. Diurnal surveys began with a 1-hr observa- tion of the colony with binoculars and spot- ting scopes from vantage points. Similar pre- dusk observation periods were also conducted. Walking surveys were made im- mediately following morning surveys. Each colony was thoroughly walked by up to 12 people simultaneously. Each person moved back and forth within a 30 m wide area and examined all prairie dog burrow openings, mounds, and adjacent areas, as well as the overall surface of the colony. Each burrow examined was marked with a footprint to as- sure complete, nonoverlapping coverage. Data recorded included: number of burrow openings (5 cm or larger in diameter); num- ber of burrow openings in "active" use, where possible to determine; number of badger excavated holes; number of plugged burrow openings; number of km walked; 'Department of Biology, Idaho State University, Pocatello, Idaho 83209. 'Biota Research and Consulting, Inc., P.O. Box 2705, Jackson, Wyoming 83001. "Department of Zoology, University of Wisconsin, Madison, Wisconsin 53706. 572 December 1982 Clark et al.: Prairie Dog Colony 573 Black -tail ^ Prairie Dog ED Gunnison's Prairie Dog 1 White -tail Prairie Dog Fig. 1- Map of the three prairie dog species' distribution along the belt transect study area (dotted line) in New Mexico, Colorado, and Utah. time spent walking; and number and species of all live vertebrates and their remains seen. Since badgers are considered the single most effective predator and directly alter a colony by digging, badger activity (percentage of burrows enlarged by badgers) was estimated on each colony (Campbell and Clark 1981). Nocturnal surveys of prairie dog colonies were conducted via spotlighting. Two spotlights were used simultaneously by each 574 Great Basin Naturalist Vol. 42, No. 4 crew of two persons. One cab-mounted light was used by the driver and the second, hand- held Ught was used by the rider seated in back. The spotlights generally allowed identi^ fication of animals out to 150 m as the truck moved at 3-8 km/hr. Travel time around a spotlight circuit varied in relation to colony size and terrain, but was usually 15-60 min- utes. On small towns (less than 10 ha) a single stationary spotlighting location was used. Spotlighting was started just after sunset and continued until around midnight and again from about 0300 hours till sunrise. All colo- nies were spotlighted for at least three con- secutive nights, but large towns were spot- lighted longer. All animals seen were identified to species and numbers recorded. To compare the two nighttime survey pe- riods for vertebrate activity and inter- observer differentials, all vertebrates seen were lumped into classes based on taxonomy and morphology (primarily body size): (1) la- gomorphs, (2) rodents, (3) flying vertebrates (birds and bats), (4) small carnivores (badgers and smaller forms), (5) large carnivores (coy- otes and bobcats), and (6) ungulates. La- gomorphs, rodents, and flying vertebrates were compared between number of species seen on the first spotlight circuit of the post- sunset and predawn survey periods. All other classes were compared on number of species seen per hour over the entire survey periods. Results Prairie Dog Colony Attributes The 47 prairie dog colonies located totaled 4760 ha and comprised 2.2 percent of the belt transect (Table 1). Black-tails occupied 0.04 percent of the transect, Gunnison colo- nies 0.2 percent, and white-tail colonies 1.9 percent. Gunnison and white-tail colonies were clumped in distribution; information was insufficient to determine if black-tail col- onies were also clumped. The first three colonies encountered were black-tails, and intercolony distances be- tween colony 1 and 2 and between 2 and 3 was 6.4 km and 86.3 km, respectively. Dis- tance to the next colony, a Gunnison town, was 355 km. Gunnison colonies fell into four distinct clumps; 127 km separated the first (N = 6 towns) and second (N = 2 towns) clumps, 30.6 km the second and third (N = 2), and 245.5 km the third and fourth (N = l). Table 1. Comparative colony characteristics among the clumps of prairie dog colonies by prairie dog species. Colony characteristics Location Number of colonies Total area (ha) Colony area (ha): Mean (SD) Range X Intercolony distance (km) Total burrow openings Burrow openings/ha: Mean (SD) Range Plugged burrows: Number and % of all openings Badger reamed: Number and % of total openings Vertebrate skeletal remains: Prairie dogs/ha Other species/ha Vegetation (cm): X Height (SD) Range Black-tails Clump 1 Clump 1 NW New Mexico Prairie Gunn- Clump 2 3 99 6 235 2 116 33(26) 10-61 46.4 2763 39(27) 3-73 2.3 5238 58(15) 47-69 3.2 1004 32.5(8.9) 23.9-41.7 209(8.4) 8.2-32.0 8.8(0.9) 8.1-9.4 106(3.8%) 13(0.2%) 4(0.3%) 102(3.4%) .366(7.8%) 65(6.8%) 0.273 0.131 0.196 0.008 0.078 0.008 64(23) 51-91 79(29) 38-112 46(22) 30-61 December 1982 Clark et al.: Prairie Dog Colony 575 The mean intercolony distance for Gunnison colonies (N= 10) was 2.4 km (SD± 1.6) range 0.5-5.3. The 33 white-tail colonies were dis- tributed in two clumps. The first group of 15 towns was about 96 km from the nearest Gimnison colony and 115 km from the sec- ond white-tail colony group (N=18). Mean intercolony distance for white-tails was 4.9 km (SD±3.0) range 0.8-11.3. Intercolony distances varied significantly interspecifically (F2,37= 17.92, P<0.01) and intraspecifically for white-tails (X2 = 56.14, df = 31, P<0.005). The interspecific size of prairie dog colo- nies varied, but insignificantly (F2,44=1.13, P>0.05). Three black-tail colonies had a mean size of 33 ha (SD±26) range 10-61, 11 Gimnison towns averaged 46 ha (SD±43) range 1.6-150, and 33 white-tail colonies av- eraged 125 ha (SD±200) range 0.2-958. A significant difference (P<0.05) in mean colo- ny size was evident within each of the three prairie dog species (black-tails X2= 40.44, df =2; Gminisons, X2 = 404.21, df = 10; white- tails, X2= 10245.69, df= 32). Plugged burrows were found in colonies of all three species. Black-tails showed 3.8 per- cent plugged of 2763 burrow openings, Gun- nisons 0.3 percent of 8987 and white-tails 1.0 percent of 85,572. The interspecific density of burrow openings per colony varied in- significantly (F2,44=1.09, P>0.10) among the three species, with black-tails showing 27.9/ha (SD±8.6, range 24-41.3), Gunnisons 17.6/ha (SD±49.2, range 8.2-179), and white-tails with 21/ha (SD±29.2 range 2.2=158). All colonies showed signs of badger activi- ty in the form of excavated holes and scats. Badger-reamed prairie dog holes (holes sus- pected of being enlarged by badger digging) on colonies varied significantly between the three prairie dog species (F2,44 = 4.67, P<0.05). Badger activity was significantly lower on black-tail colonies than on Gun- nison (t = -7.42, df=12, P<0.01) and white- tail colonies (t = -8.69, df = 34, P<0.01), but was not significantly different on Gunnison and white-tail colonies (t = -1.88, df = 42, P>0.10). Badger activity was positively and significantly correlated (P<0.01) to colony size and to the number of burrow openings on each colony for both Gunnisons (r = 0.8729, t = 5.37 and r = 0.9431, t = 8.51, respectively) and white-tails (r = 0.9084,' t = 12.10 and r = 0.9845, t = 31.25, respective- ly), but not for black-tails, nor were there Table 1 continued. dog species ison s White-tails Clump 3 Clump 4 Total Clump 1 Clump 2 Total SW Colorado EUtah - EUtah _ 2 10 1 150 11 511 15 566 18 3584 33 4150 5(4) 2-8 1.6 668 150 2077 33(26) 2-73 2.4 8987 38(37) 9-121 5.5 8993 199(249) 0.2-958 4.4 76,579 125(200) 0.2-958 4.9 85,572 96.2(69.1) 47.3-145 13.8(-) 31.7(39.4) 8.1-145 19.8(10.3) 2.,3-40.5 30.8(37.6) 5.1-160 25.8(28.8) 2.3-160 0 8(0.4%) 25(0.3%) 23(0.3%) 40(0.3%) 790(0.9%) 50(7.2%) 235(11.3%) 716(7.8%) 769(9.3%) 16,469(17.5%) 17,238(13.8%) 0.90 0.30 0.287 0.012 0.209 0.018 0.150 0.0179 0.159 0.0209 0.157 0.020 84(32) 61-107 61 673(28) 30-112 39(23) 0-91 46(15) 15-71 42.6(19) 0-91 576 Great Basin Naturalist Vol. 42, No. 4 40 SPCXIES z « Bt«k-T«il • QhWWiMW • WMtv-Tol 0) y 30 o UJ CL • • U. 9 o o • • o q: « UJ • n • ••o • •M or O 2 • 13 5 • Z »o • SPEARMAN RANK CORRELATION r«0.8l 20 40 60 80 KX) 120 »40 160 180 >200 PRAIRIE DOG COLONY SIZE (ha) Fig. 2. Relationship between black-tail, Gunnison, and white-tail prairie dog colony size and number of vertebrate species observed on each colony. significant correlations with burrow density or intercolony distances. Associated Vertebrate Species A total of 107 species and subspecies, ver- tebrate animals, including one amphibian, 25 reptiles, 51 birds, and 30 mammals, were ob- served on prairie dog colonies (Table 2). A larger number of vertebrate species were seen on white-tailed colonies than on colonies of the other two prairie dog species; 88 per- cent of the surface area of all prairie dog col- onies was in white-tail colonies. Six species of mammals, 7 species of birds, 2 species of reptiles, and no species of am- phibians were common to colonies of all three prairie dog species. In contrast, 7 mam- mal, 11 bird, 8 reptile, and no amphibian species were common to colonies of two prairie dog species, and 17 mammal, 33 bird, 15 reptile, and one amphibian species were present on colonies of only one prairie dog species. The relationship between the number of vertebrate species seen on prairie dog colo- nies of varying sizes is shown in Figure 2. A Spearman Rank Correlation (r = .81) showed that larger towns contained more vertebrate diversity than smaller colonies. Four rattlesnake species and subspecies were found, 1 western diamondback, 4 Hopi, 7 prairie, and 6 midget-faded rattlesnakes. Rattlesnakes on black-tail colonies occurred at 0.02/ha, on Gunnison at 0.02/ha, and on white-tail at 0.002/ha. The Hopi were on a single 148 ha Gunnison colony. Eleven raptor species, including one eagle, 4 hawks, 3 falcons, and 3 owls, were seen. Burrowing owls (N = 99) occurred on 19 towns and at a density of 0.04 owls/ha. The greatest density was 15 owls on a 10 ha town. December 1982 Clark et al.: Prairie Dog Colony 577 CoTra^C^and'utl''*' '^''''' '""^ '"''''''"'■' ^''"'"^ °" '^' '°'°"^'' °^ '^'"'' P^^'"^ ^°g ^P^^''^^ '" New Mexico, Vertebrate species and subspecies Prairie dog species Black-tail Amphibians Great Basin spadefoot toad (Scophiopus intermontanus) Totals Reptiles Mountain short-homed lizard {Phnjnosoma douglassi hernandesi) Eastern short-horned lizard (P. douglassi brevirostre) Desert short-homed lizard {P. douglassi ornatissiimim) Texas horned lizard (P. cornutum) Sagebmsh lizard {Sceloponis gra.siosiis) Northem plateau lizard (S. undulatits elongatus) Northern whiptail (Cnemidophorus tigris septentrionalis) Westem whiptail (C. tigris) Little striped whiptail (C. inornatus) Chihuahua whiptail (C. exsanguis) Checkered whiptail (C. tesselattis) Leopard lizard {Crotaphytus wislizenii) Lesser earless lizard (Holbrookia maculata) Side-blotched lizard {Vta stanshuriana) Westem coachwhip {Masticophis flagellum testaceus) Great Basin gopher snake [Pituophis melanoleucus deserticola) Bullsnake {P. m. saiji) Utah milk snake {Lampropeltis triangulum taylori) Wandering garter snake (Tfiamnophis elegans vagrans) Painted desert glossy snake {Arizona elegans philipi) Westem diamondback rattlesnake (Crotahis atrox) Midget faded rattlesnake (C. viridis concolor) Prairie rattlesnake (C. v. viridis) Hopi rattlesnake (C. v. nuntius) Westem box turtle {Terrapene ornata) Totals (n = 25) Birds Canada Goose (Branta canadensis) Mallard (Anas platyrhynchos) Green-winged Teal (Anas crecca) Turkey Vulture {Cathartes aura) Marsh Hawk (Circus cyaneus) Red-tailed Hawk (Buteo jamaicensis) Fermginous Hawk (B. regalis) Swainson's Hawk (B. swainsoni) Golden Eagle (Aquila chrysaetos) Prairie Falcon (Falco mexicanus) Merlin (f. columbarius) Kestrel (F. sparveriiis) Short-eared Owl (Asio flammeus) Great Homed Owl (Bitbo virginiamis) Burrowing Owl {Athere cunicularia) Sage Grouse (Centrocercus urophasianus) Gambel's Quail (Lophortyx gambelii) Killdeer (Charadrius vociferus) Westem Sandpiper {Calidris rnauri) Mouming Dove (Zenaida macroura) Common Nighthawk (Chordeiles minor) Lesser Nighthawk (C. acutipennis) Poor-will (Phalaenoptilus nuttallii) Gunnison X X X X X X X X X 16 White-tail 15 X X X X X X X X X X X X X X X X X X X X X 578 Great Basin Naturalist Vol. 42, No. 4 Table 2 continued. Vertebrate species and subspecies Prairie dog species Black-tail Gunnison White-tail Birds continued. Western Kingbird (Tyrannus verticalis) Say's Phoebe (Sayornis saya) Gray Flycatcher (Empidonax wrightii) Ash-throated Flycatcher {Myiarchus cinerascens) Horned Lark {Eremophila alpestris) Violet-green Swallow (Tachycineta thalassina) Barn Swallow (Hirundo rustica) Rough-winged Swallow {Stelgidopteryx ruficollis) Pinyon Jay {Gymnorhinus cyanocephahis) Black-billed Magpie {Pica pica) Common Raven (Corvus corax) Rock Wren {Salpinctes olsoletus) Sage Thrasher {Oreoscoptes montanus) Curve-billed Thrasher {Toxostoina curvirostre) Mockingbird (Mimus polyglottos) Mountain Bluebird {Sialis currucoides) Northern Shrike {Lanius excubitor) Loggerhead Shrike (L. ludovicianus) Western Meadowlark {Strunella neglecta) Eastern Meadowlark (S. magna) Brewer's Blackbird {Euphagus cyanocephahis) Western Tanager (Piranga lucoviciana) Grasshopper Sparrow {Ammodramus savannariim) Vesper Sparrow (Pooecetes gramineus) Black-throated Sparrow {Amphispiza bilineata) Sage Sparrow (A. belli) Lark Sparrow (Chondestes grammacus) Brewer's Sparrow (Spizella breweri) Totals (n = 51) Mammals Bat (unidentified) Desert cottontail {Sylvilagus auduhoni) Mountain cottontail (S. nuttalli) White-tailed jackrabbit (Lepus townsendi) Black-tailed jackrabbit (L. californicus) Thirteen-lined ground squirrel {Spermophilus tridecemlineatiis) Wyoming ground squirrel (S. elegans) Whitetail antelope squirrel {Ammospermophilus leucurus) Pocket gopher (Tfiomomys sp.) Valley pocket gopher (T. bottae) Plains pocket gopher (Geoniys bursariiis) Least chipmunk {Eutamius mini7nus) Woodrat (Neotoma sp.) Bushytail woodrat [N. cinerea) Southern plains woodrat (N. micropus) Ord kangaroo rat (Dipodomys ordi) Bannertail kangaroo rat (D. spectabilis) Deer mouse {Peromyscus maniculatus) Vole (Microtus sp.) Muskrat {Ondatra zibethica) Coyote {Canis latrans) Swift fox {Vulpes velox) Domestic dog (Canis familiarus) Domestic cat {Felis domesticus) Long-tailed weasel {Mustela frenata) 23 X X X X X X X X X X X X X X X X X X X X X X X 44 December 1982 Table 2 continued. Clark ET AL. Prairie Dog Colony 579 Vertebrate species Prairie dog species and subspecies Black-tail Gunnison White-tail Mammals continued. Short-tailed weasel (A/, erminea) Badger (Taxidea taxus) Striped skunk (Mephitis mephitis) Mule deer [Odocoileus hemionus) Pronghom [Antilocapra americana) X X X X X X X X X X Totals (n = 30) 10 16 23 Only two live weasels were found. a Discussion bridled weasel in New Mexico (61 ha town) and a short-tailed weasel in Utah (102 ha town). Both were sighted in daylight. Fourteen live badgers were seen on 10 col- onies (1 badger/ 183 ha, range 0.2-522). All badgers were the sole mustelid species seen on its respective town except for 5 badgers on a 946 ha town (1 badger/ 189 ha). All towns showed signs of badger activity, al- though varying greatly in density. Coyotes (N = 29) were seen on 18 towns (1 coyote/ 115 ha). An additional 12 towns had coyote scats. Kit foxes (N = 5) were seen on 4 tovms that had a mean size of 62 ha (range 20-148). Two foxes were on a 26 ha town. Numerous vertebrate remains were found on prairie dog towns (Table 1). In all, 1597 remains of individual prairie dogs and 202 other vertebrate remains of at least 16 spe- cies were found. Prairie dog remains oc- curred at about one skeleton/5 ha and re- mains of other vertebrates at about 1/50 ha. Nocturnal survey results showed that some vertebrates had a differential observability between the postsunset and predawn survey periods. The front and back spotlight per- sonnel observed species and categories of ver- tebrates differentially. Three categories of vertebrate sightings-flying forms (X2=159, df=l, P<0.01), ungulates (X2=ll, df=l, P<0.01), and large carnivores (X2=10.5, df=l, P<0.05)— showed significant differ- ences in observability, which may reflect real differences in activity, with most being seen per unit effort in postsunset periods. Rodents (X2= 18.8, df = 1, P<0.01) and fly- ing vertebrates (X2 = 45.1, df=l, P<0.01) were sighted significantly more by the driver than the rider in back. The driver and rider saw no significant differences between all other classes. Prairie dog colonies occupied only a small portion of the survey area (2.2 percent); less than 1 percent for black-tail and for Gun- nison and about 1.9 percent for white-tail. The U.S. Forest Service (1981) found black- tails on Thunder Basin National Grassland, Wyoming, to occupy 1.3 percent. Elsewhere in Wyoming, Campbell and Clark (1981) found black-tails to occupy only 0.7 percent of a 1036 km2 and white-tails 3.2 percent of 336 km2. An area in southwest Wyoming contained 63 white-tail towns and occupied 25 percent of the 259 km2 area (Clark and Campbell, unpubl. ms.). White-tails in two other areas in southern Wyoming occupied 7.2 percent and 8.9 percent of the study area (Martin and Schroeder 1979, 1980) (Table 3). In comparing the three black-tail colonies in our study with 186 other black-tail towns elsewhere, our 11 Gunnisons with one other, and our 33 white-tail towns with 354 others (Table 3), all the towns we surveyed fall within the ranges of colony sizes and burrow opening features previously reported. Our survey of Gunnison prairie dogs appears to be the first relatively large sample. We found a mean intercolony distance of 46.4 km for black-tails, 2.4 km for Gunnisons and 4.9 km for white-tails, whereas Campbell and Clark (1981) found 4.7 km (SD±2.7) range 1.6-11.3 for white-tails in two large rela- tively undisturbed areas in Wyoming. Prairie dog colonies were found clumped in suitable habitat, and nearby colonies served as sources for colonizing animals. White-tail colonies (N = 19) in the Vernal, Utah, clump were part of a much larger com- plex of colonies estimated to be at least 5,000 ha. No comparable situation for the black-tail or Gunnison was found. 580 Great Basin Naturalist Vol. 42, No. 4 Prairie dogs are known to plug burrows in response to predator investigations, death of prairie dogs, and other disturbances (Koford 1958, Henderson et al. 1969). We found black-tails plugged 3.8 percent of their bur- row openings, Gunnisons 0.2 percent and white-tails 0.9 percent. In comparison, Campbell and Clark (1981) found 0.0005 per- cent plugged burrow openings for black-tails and 0 percent for white-tails in Wyoming. Gold (1976), Bonham and Lerwick (1976), and Hansen and Gold (1977) noted that black-tail prairie dogs manipulate soil and in- crease plant and animal density and therefore may be viewed as ecosystem regulators. Uresk and Bjugstad (in press) noted that peak plant production of aboveground herbage over their five-year study occurred where prairie dogs only grazed during the last four years of the study, rather than under the other comparative treatments (prairie dogs and steers, steers only, neither). Our surveys found 107 vertebrate species and subspecies on or over prairie dog colo- nies. Additional species occupying prairie dog colonies were reported by Tyler (1968), Martin and Schroeder (1979, 1980), and Campbell and Clark (1981). Collectively, over 140 vertebrate species have been re- ported associated with prairie dogs. A general account of many of the more conspicuous vertebrates and their inter- relationships with prairie dogs was discussed by Koford (1958). Prairie dogs improve habi- tat for prairie animals that are benefited by holes, unvegetated areas, and short vegeta- tion and for those that feed on prairie dogs. The prairie dog burrow is a critical element in prairie dog survival and allows them to es- cape the extremes of temperature, for ex- ample, and benefits numerous other verte- brates as well (Stromberg 1978). Desert cottontails, burrowing owls, swift foxes, rattlesnakes, and some species of plants are enhanced by prairie dog activities (Uresk and Bjugstad, in press). Sixty-three percent more small mammals, other than prairie dogs, were live-trapped on pastures used by steers and prairie dogs than on steer-only pastures (O'Melia 1980). O'Melia also found prairie dogs significantly decrease arthropod populations. Table 3. Prairie dog colony characteristics for black-tail, white-tail, and Gunnison prairie dog species found in this and other studies conducted in Wyoming, Colorado, Montana, Kansas, and South Dakota. Prairie dog species and location Number of colonies Colony area (ha) Total X (SD) range Black-tail Prairie Dog Southeast New Mexico Central Wyoming Eastern Wyoming Central South Dakota Western South Dakota Northern Colorado Great Plains Central Kansas Northern Wyoming Gunnison Prairie Dog Northeast New Mexico Southwest Colorado White-tail Prairie Dog Eastern Utah South central Wyoming Northwest Wyoming Southwest Wyoming South central Wyoming Southern Wyoming Southern Wyoming Southern Montana North central Colorado 3 21 2 151 2 1 1 7 11 1 33 25 4 63 1 164 81 15 1 99 33(26)10-61 731 35(44)1-189 123 66(— )26-97 1,283 8.5(— )— 47 580 511 6 13 4,006 60,665 285 9 3(-)- 47(-)- 83(— )2.8-359 33(26)2-73 6(-)- 4,150 125(200)0.2-958 1,085 43(46)2-184 3,055 764(— )121-1416 3,992 63(— )0.4-671 13(-)- 24(— )0.8-510 54(— )0.4-414 19(-) 9(-)- December 1982 Clark et al.: Prairie Dog Colony 581 Prairie dogs provide food to the black- footed ferret (Hillman and Clark 1980), bad- gers, foxes, coyotes, bobcats, and weasels as well as to Golden Eagles, Ferruginous Hawks, and Swainson's Hawks (Campbell and Clark 1981). The U.S. Bureau of Land Management (1979) noted that prairie dog towns also provide nest sites for Moimtain Plovers {Charadriiis montanus) and McCown's Longspur (Calcariiis mccownii) and benefit other birds such as Killdeer, East- em Kingbirds {Tijrannus tyranniis). Upland Sandpiper, Long-billed Curlews {Numenius americamis) and Mourning Doves, to men- tion a few. Sharp-tailed (Pedioecetes phasia- nollns) and Sage grouse sometimes strut on prairie dog towns (McEneany and Jensen 1974). A large number of reptiles use prairie dog holes for thermoregulation and pro- tection from predators. Reptiles eat arthro- pods inhabiting both burrows and the surface of the colony (Wilcomb 1954, Clark 1977). Evidence of human presence— roads, spent shell casings, plowing, and some evidence of poisoning— was obvious on many colonies. Our observations in this study and elsewhere indicate that most prairie dog colonies are negatively influenced by humans; only a few areas still contain large, relatively undis- turbed colonies. This brief account of prairie dog colony at- tributes and the relationships of some associ- ated vertebrate species with prairie dogs shows that numerous benefits may be ac- crued to some of those vertebrates. Prairie dog colonies may constitute peaks in species diversity and biomass for small vertebrates found nowhere else in the prairie ecosystem. Needed are precise community ecology stud- ies, with adequate controls near prairie dog towns, to quantify relative degrees of associ- ation between the various vertebrate species and prairie dogs and to identify the types of relationships that exist. Acknowledgments Many people provided direct aid to this project. James Leiber and Gene Bell of Mid- America Pipeline Co., Inc., Tulsa, Oklahoma, provided the opportunity and support for the survey. Dana Shuford, U.S. Bureau of Land Table 3 continued. Burrow openings Percent Percent badger reamed — Density/ha Total X (SD) Range plugged Sources 2,763 33(9)24-42 3.8 3.4 This study 17,095 21(24)11-67 0.0 10.0 Campbell & Clark 1981 6,015 49(10)38-52 0.9 2.6 Clark & Campbell unpubl. ~ — (— )— — — Linder et al. 1972 - 135(— )131-140 — _ King 1955 289 96(-)- — — Tileston & Lechleitner 1966 — 5(— )15-67 — — Koford 1958 6,344 135(-)- — — Smith 1958 29,215 50(-)~ 2.2 - Martin & Schroeder 1980 8,987 32(39)8-145 2.8 7.8 This study 321 57(-)- — — Fitzgerald & Lechleitner 1974 85,572 26(29)2-160 0.9 13.8 This study 27,779 25(26)9-129 0.0 27 Campbell & Clark 1981 6,755 2.2(— )l-6 — _ Clark et al. unpubl. 168,761 42(— )0.7-21.3 — 24.7 Clark & Campbell unpubl. and Martin & Schroeder 1979, 1980 827 64(-)- — _ Clark 1977 105,497 26(— )0.8-41 - — Martin & Schroeder 1979 129,969 32(-)- 4.3 — Martin & Schroeder 1980 — -(-)- — — Flath 1979 252 28(-)- - - Tileston & Lechleitner 1966 582 Great Basin Naturalist Vol. 42, No. 4 Management, helped facilitate the project. Janice Hutton, David Hayden, and Ron Free- man of Woodward/ Clyde Consultants, San Francisco, California, provided adminis- trative and field assistance. In addition to ourselves, survey teams con- sisted of Marcia Casey, Janette Johnson, Che- ryl Lorenz, Suzanne Martell, Ronald Pace, Arme Rathbun, Sue Wandersee, JoAnn Cam- enzind, Marilyn McElheney, Kane Bright- man, John Hoak, and Karen Jerger. Their un- tiring efforts, keen observations, and total commitment to a quality survey were in- valuable. We would like to thank Jeff Marks for aiding with bird names. We sincerely thank all those people who directly aided in the survey and many others not mentioned by name who helped indirectly. Literature Cited BoNHAM, D. C, AND A. Lerwick. 1976. Vegetation changes induced by prairie dogs on shortgrass range. J. Range Manage. 27:221-225. Campbell, T. M., and T. W. Clark. 1981. Colony char- acteristics and vertebrate associates of white- tailed and black-tailed prairie dogs in Wyoming. Am. Midi. Nat. 105:269-276. Clark, T. W. 1977. Ecology and ethology of the white- tailed prairie dog (Cijnomys leucurus). Mil- waukee Public Mus. Publ. Biol. Geol. No. 3. 97 pp. Fitzgerald, J. P., and R. R. Lechleitner. 1974. Obser- vations on the biology of Gunnison's prairie dog in central Colorado. Am. Midi. Nat. 92:146-163. Flath, D. L. 1979. Status of the white-tailed prairie dog in Montana. Proc. Mont. Acad. Sci. 38:63-67. Gold, I. K. 1976. Effects of blacktail prairie dog mounds on shortgrass vegetation. Unpublished thesis. Colorado State Univ., Fort Collins, Colorado. 39 pp. Hansen, R. M., and I. K. Gold. 1977. Blacktail prairie dogs, desert cottontails, and cattle trophic rela- tions on shortgrass range. J. Range Manage. 30:210-214. Henderson, F. R., P. F. Springer, and R. Adrian. 1969. The black-footed ferret in South Dakota. South Dakota Dept. Game, Fish, and Parks Tech. Bull. 4. 37 pp. King, J. A. 1955. Social behavior, social organization, and population dynamics in a black-tailed prairie dog town in the Black Hills of South Dakota. Univ. Michigan Contrib. Lab. Vert. Biol. 67:1-123. KoFORD, C. B. 1958. Prairie dogs, whitefaces, and blue grama. Wildl. Monogr. No. 3. 78 pp. Linder, R. L., R. B. Dahlgren, and C. N. Hillman. 1972. Black-footed ferret-prairie dog inter- relationships. Pages 22-37 in Symp. Rare and En- dangered Wildl. of the SW U.S., Sept. 22-23, 1972. Albuquerque, New Mexico. Martin, S. J., and M. H. Schroeder. 1979. Black-footed ferret surveys on seven coal occurrence areas in southwestern and southcentral Wyoming, June 8 to September 25, 1978. Final report, Wyoming State Office BLM. 37 pp. 1980. Black-footed ferret .surveys on seven coal occurrence areas in Wyoming, Febru- ary-September, 1969. Final report, Wyoming State Office BLM. 39 pp. McEneany, T. p., and J. L. Jensen. 1974. Status of the black-tailed prairie dog on the Charles M. Russell National Wildlife Refuge. USFWS. Lewiston, Montana. 7 pp. O'Melia, M. E. 1980. Competition between prairie dogs and beef cattle for range forage. Unpublished thesis. Oklahoma State Univ., Stillwater, Okla- homa. 33 pp. Pizzimenti, J. J. 1981. Increasing sexual dimorphism in prairie dogs: evidence for changes during the past century. Southw. Nat. 26:43-47. Smith, R. E. 1958. Natural history of the prairie dog in Kansas. Univ. Kans. Mus. Nat. Hist, and St. Biol. Surv. Misc. Publ, 16:l-.36. Stromberg, M. R. 1978. Subsurface burrow connections and entrance spatial pattern of prairie dogs. Southw. Nat. 23:173-180. TiLESTON, J. v., AND R. R. Lechleitner. 1966. Some comparisons of the black-tailed and white-tailed prairie dog in north central Colorado. Am. Midi. Nat. 75:292-316. Tyler, J. D. 1968. Distribution and vertebrate associates of the black-tailed prairie dog in Oklahoma. Un- published dissertation. Univ. of Oklahoma, Nor- man, Oklahoma. 85 pp. WiLCOMB, M. J. 1954. A study of prairie dog burrow sys- tems and the ecology of their arthropod in- habitants in central Oklahoma. Unpublished dis- sertation. Univ. of Oklahoma, Norman, Oklahoma. 154 pp. Uresk, D. W., and a. J. BjuGSTAD. Effects of prairie dogs and cattle on the vegetation of the northern high plains. Proc. 7th No. Amer. Prairie Conf. (in press). U.S. Bureau of Land Management. 1979. Habitat man- agement plan-prairie dog ecosystem. Montana State Office, Billings, Montana. 61 pp. U.S. Forest Service. 1981. Thunder Basin prairie dog management. USDA. Environmental Assessment. Thunder Basin Nat. Grasslands, 809 South 9th St., Douglas, Wyoming 82633. DISTRIBUTION OF THE MOSS FAMILY GRIMMIACEAE IN NEVADA Matt Lavin' .Abstract.- Twenty-six taxa of Grimniiaceae are listed from Nevada, all representing the genera Grimmia and Rhacomitnum. Rhacomitriwn heterostichum (Hedw.) Brid. var. heterostichum and Grimmia atricha C. Muell & Kindb. ex Mac. & Kindb. are listed for the first time as occurring in Nevada. Within the state, the Mohave Desert the Great Basin desert, and the Sierra Nevada display unique composition of members of the Grimmiaceae Grimmia anochn is the most widespread moss in the state. Others, such as Grimmia rivulare, G. conferta, and G. alpicola in- habit only the montane environments of northeastern Nevada. Nearly 1000 collection or observation sites throughout Nevada were visited during the past four years that the moss family Grim- miaceae has been under investigation. Two genera make up this family in Nevada, Grimmia and Rhacomitrium. Schistidium, in this paper, is used as a subgenus under the genus Grimrnia. These mosses constitute a modest percent- age of biomass in many plant communities of Nevada. They are restricted to rock habitat, although Grimmia occidentalis is occasionally foimd on the base of trees near stream sides. Grimmia subgenus Grimmia generally occurs on dry, exposed rock, but Griminia subgenus Schistidium, and Rhacomitrium occur on rock that is or has been inundated by water from spring snow melt, or deep within rock crevices that offer protection from exposure to the sun and heat. Members of the Grimmiaceae display unique composition in three geographical areas in Nevada. These areas are (1) the Mo- have Desert in the very southern part of the state, (2) the Great Basin desert and associ- ated mountain ranges, which includes most of the state, and (3) the Sierra Nevada in the very western portion of the state. The Mohave Desert vegetation is domi- nated by such vascular plants as Yucca hrevi- folia (Joshua tree) and Larrea tridentata (creosote bush). In this desert, Grimmia or- bicularis, G. wrightii, G. anodon, and G. af- finis are the dominant and practically the only mosses, sometimes codominating the cryptogamic flora with Tortula inermis, Cros- sidium ahherans, C. griseum, and the hepatic Targonia heterophylla. Grimmia af finis, orig- inally described as autoicous, is dioicous in Nevada. This agrees with Flowers (1973). Mesic habitats in this southern desert are found in unusual abundance in the deep can- yons of the Spring Mountains just west of Las Vegas. Here, Grimmia ovalis, G. pulvinata, G. stricta, and G. atricha grow, along with other common mosses such as Anacolia men- ziesii, Brachythecium collinum, Encalypta in- termedia (i.e. Encalypta intermedia), and Or- thotrichum cupulatum. Grimmia ovalis forms unusually long stems in this area, up to 6 cm in length. G. pulvinata, G. atricha, and G. stricta oc- cur throughout the Pacific Northwest and might, therefore, be expected to occur in more northerly portions of Nevada. How- ever, the only Nevada collections come from the southern part of the state. Grimmia at- richa is reported from the Spring Mountains by a collection of Dr. H. Mozingo, University of Nevada, Reno. This represents possibly the most southern distribution for this species. Many endemic and relictual vascular plants occur on this range and this moss could be a holdover from Pleistocene vegetation. The Great Basin Desert vegetation is domi- nated by Artemisia tridentata (big sage), Pinus monophylla (single leaf pinyon pine), and Juniperus osteosperma (Utah juniper). Cryptogamically, it is dominated by Grimmia tenerrima, G. anodon, and G. calyptrata. These three mosses commonly grow together at practically all elevations. Grimmia calyp- trata inhabits predominantly the north- 'P.O. Box 13494, Reno, Nevada 89507. 583 584 Great Basin Naturalist Vol. 42, No. 4 western exposures, and G. tenerrima and G. anodon assume the somewhat more protected northeast exposures. As G. tenerrima is dioicous, the male and female plants appear as separate entities due to differences in the lengths of the hairpoints. The male plant, with very short hairpoints, is often confused with G. anodon. G. anodon, however, is au- toicous, thereby making the two easily separable. Grimmia anodon has the most widespread distribution of probably any plant species in Nevada. Tlie plants in southern Nevada have a much greener appearance, shorter hair- points, and more erect stems than the same species from the north. Grimmia calyptrata, although abundant in northern Nevada, is not foimd in southern Nevada. It is, apparently. Table 1. A list of the Grimmiaceae in Nevada. No- menclature for Grimmia subgenus Grimmia and Rhcico- mitriwn generally follows Lawton (1971). Nomenclature for the specific epithet in Grimmia subgenus Schistidium is based on Deguchi (1979). Grimmia (subgenus Grimmia) af finis Hornsch. anodon B.S.G. calyptrata Hook, ex Drumm. laevigata Brid. montana B.S.G. orbicularis Bnich. ovalis (Hedw.) Lindb. plagiopodia Hedw. poecilostoma Card. & Seb. pulvinata (Hedw.) Small tenerrima Ren. & Card. (SY = G. alpestris [Web. & Mohr] Nees) torquata Hornsch. var. torquata trichopbijlla Grev. wrightii (Sull.) Aust. Grimmia (subgenus Schistidium) alpicola Hedw. (SY = G. agassizii (Sull. & Lesq. ex Sull.) Jaeg. & Sauerb. See Bremer (1980) and Deguchi (1979 and 1979a) for nomenclature of this entity.) alpicola var. latifolia (Zett.) Moll. ambigiia Sull. apocarpa Hedw. atricha C. Muell & Kindberg ex Macoun & Kindberg conferta Funck flaccida (DeNot.) Lindb. occidentalis Lawton pacifica Lawton rivularis Brid. (SY = G. alpicola var. rivularis [Brid.] Wahlenb.) stricta Turn. Rhacomitrium beterostichum (Hedw.) Brid. var. beterosticbum (verified by Lawton, 1981) ecologically replaced by G. orbicularis in the south. They both inhabit very exposed rock and .superficially resemble each other with regard to long hairpoints on the leaves and the large rounded clumps they both form (re- sembling small hedgehogs). On the granitic boulders that follow Salmon Falls Creek in Elko County of north- eastern Nevada, G. poecilostoma occurs, far north of its otherwise reported range. This moss has supposedly a more southern distri- bution, including New Mexico, Texas, and Arizona. In extremely dry situations, such as the flat desert country in westcentral Nevada, Grimmia anodon is almost the only existing moss, inhabiting mostly low lying, flat rock. Mosses commonly occurring with the Grimmia species of this part of the Great Ba- sin include Brachythecium collinum, Dicrano- wesia crispula, Encahjpta intermedia, Or- thotrichum cupulatiim, O. jamesiamim, O. laevigatum f. macounii, Pseudoleskeella tecto- rum, Pterygoneiirnm ovatiim, P. subsessile, Timmia megapolitana, Tortula papillosissima, and T. ruralis. Following spring snow melt rvuioff in the high mountains of the Great Basin, rocks are inhabited by a few members of Grimmia sub- genus Schistidium. These species include G. occidentalis, G. rivtdare, G. alpicola, and G. pacifica. These mostly occur by themselves, but may occur with such mosses as Lescuraea incurvata or Orthotrichum rividare. These mosses nearly always inhabit montane to al- pine environments. However, G. occidentalis, along with Orthotrichum rividare, was ob- served to occur in the pinyon-juniper wood- lands of the Virginia Range of westcentral Nevada. Grimmia pacifica, collected in the Santa Rosa Range of northcentral Nevada, is a very interesting plant. The spores measure up to 30 jum in diameter, and the upper portions of the leaves are keeled with some of the lower leaf margins slightly recurved. This lends some doubt as to its identification, but Law- ton (1980) indicates that this species is rarely collected and it may be another variable spe- cies in the Grimmiaceae. Personal observa- tion of this specimen plus other specimens from the Pacific Northwest have suggested December 1982 Lavin: Nevada Mosses 585 that this entity may be nothing more than an ecotype of G. apocarpa. The most successful member of Schisti- dium in Nevada is G. flaccida. It is the only Schistidium that inhabits extremely dry rock outcrops in this state. It is mostly not found on extreme exposures (southwest faces) but is more commonly found within protected rock crevices on northern exposures. In Nevada, it is interesting to note that the most common Grimmia, G. anodon, and the most common Schistidium, G. flaccida, are the only two members of the Grimmiaceae in Nevada that completely lack a peristome. The latter spe- cies is common in northern Nevada even though records appear to the contrary. In the northeastern portion of Nevada, the Great Basin has a greater diversity of dry rock Grimmia. Aside from the three domi- nants previously listed, G. ovalis, G. poecilo- stoma, G. apocarpa, and G. flaccida occur in greater abimdance. Grimmia ambigua and G. conferta, also occur in this area, but more commonly inhabit deep crevices or shady north faces of rocks and boulders. Grimmia conferta, in its habitat and general appear- ance, appears to be a link that connects the subgenera Schistidium and Grimmia. In Wyoming, Idaho, and parts of northeastern Nevada, this moss forms small rounded clumps on exposed rock surfaces, a habit typ- ical of members of Grimmia subgenus Grimmia. The Jarbidge, Ruby, East Humboldt, and Snake mountains are located in the north- eastern portion of Nevada. It is in these moimtains that many vascular plants from the Rocky Mountains and Pacific Northwest make their only appearance in Nevada. These vascular plants include Abies lasio- carpa, Silene acaulis, Saxifraga caespitosa, Arctostaplujlos iwa-ursi. Primula parryi, Se- laginella selaginoides, and Astragalus aborigi- num. Along the same lines, the montane en- vironments in northeastern Nevada provide habitat for G. alpicola, G. ambigua, G. con- ferta, G. rivulare, and G. alpicola var. latifolia. The Sierra Nevada, characterized by the Jeffery pine, lodgepole pine, red fir, and whitebark pine forests, is dominated, with re- gard to mosses, by Grimmia tenerrima and G. montana. Grimmia trichophylla and G. tor- quata are present and appear to be imique. in Nevada, to this area. Of particular interest in the Sierra Nevada is the absence of G. anodon. This is unusual because this moss is the most widespread in the state, occurring in all counties and elevations from below 300 m in the south to over 4000 m on top of Mount Moriah in east central Nevada. One specimen of G. anodon was found in the Sierra Nevada just west of Carson City on Snow Valley Peak. It is a very atypical speci- men in that its seta ranges from straight to arcuate and the calyptra is large and cucul- late, as well as typically small and mitrate. Substrate does not appear to have a role in the exclusion of G. anodon from the Sierra Nevada. The granodiorites of the Fox, Sell- inite. Granite, and Wassuk ranges of western Nevada are inhabited frequently by G. ano- don. These granodiorites are similar and gen- etically related to the granodiorite of the Sierra Nevada Batholith (Hibbard 1982). Snow pack or summer aridity of the Sierra doesn't play a role either: G. anodon occurs in high montane habitats throughout the Great Basin, as well as throughout the desert areas of the state. Rock inundated by spring snow melt runoff in the Sierra Nevada provides habitat for both Grimmia occidentalis and G. apocarpa. These are fairly common mosses throughout the entirety of the high montane habitats in the southern Sierra. Also present is Rhacom- trium heterostichum var. heterostickum, but very rarely in the Nevada portion of the Sierra Nevada. These wet rock mosses occur by themselves and dry rock Grimmia occur commonly with Dicranoweisia crispula, En- clypta vtdgaris var. vulgaris, Homalothecium nevademe, Orthotrichum laevigatum, f. ma- counii, O. praemorsum, O. pylaissi, Tortula papillosissima, and T. princeps. Acknowledgments This study was funded in part by a grant from Sigma Xi. I am especially grateful to Dr. Elva Lawton for determination and veri- fication of many specimens. I thank Dr. Rich- ard Rust and Dr. H. Mozingo, University of Nevada, for providing much of the field transportation throughout Nevada. Addition- ally, I thank Dr. Dale Vitt, University of Al- berta, for identifying the associated species, namely Orthotrichum. 586 Great Basin Naturalist Vol. 42, No. 4 G. af finis G, anodon G, calyptrata G. laevigata G. montana G. orbicularis G. ovalis G. T3la;j:iQPodia G, poecilostoma G. pulvinata G. tenerrima G. torquata Fig. 1. Distribution of the Grimmiaceae of Nevada. Representative specimens are deposited at the University of Washington Seattle (WTU). Distribution data come from collections and observations of the author, collections of Dr. H. Mozingo, the University of Nevada, and literature citations given by Lawton (1958). Grimmia torquata was not found during the course of this study so the location given by Lawton (1958) is used. December 1982 Lavin: Nevada Mosses 587 G, trichophvlla \i f^. wri^^htii G, alpicola var, alpicola G, _alpicola ^ar. latifolia G. ambigua G. apocarpa G, atricha G, conf ert G, flaccida G. occidental j r \ \ G. pacifica G. rivularis Fig. 1 continued. 588 Great Basin Naturalist Vol. 42, No. 4 G, striata R, heterostichunr var. hetorostichunf Fig. 1 continued. Literature Cited Bremer, B. 1980. Proposal to reject the names Grimmia alpicola Sw. ex Hedw. and Schistidiinn alpicola (Sw. ex Hedw.) Linipr. (Griminiaceae). Taxon 29:337-339. Deguchi, H. 1979. A revision of the genera Grimmia. Schistidium, and Coscinodon (Musci) ol Japan. J. Sci. Hiroshima University, Series B, Div. 2, 16:121-2.50. 1979a. Les veritables caracteres de Schistidium alpicolum (Sw. ex Hedw.) Limpr. et son nouveau synonyme Schistidium agassizii Sull. et Lesq. Rev. Bryol. Lichenol. 45(4):425-435. Flowers, S. 1973. Mosses of Utah and the West. Brig- ham Young University Press, Provo, Utah. HiBBARD, M. 1982. Personal communication. University of Nevada, Reno. Lawton, E. 1958. Mosses of Nevada. Brvologist 61:314-340. 1971. Moss Flora of the Pacific Northwest. Hat- tori Botanical Laboratory. Nichinan. 1980. Personal communication. University of Washington, Seattle. INSULAR BIOGEOGRAPHY OF MAMMALS IN THE GREAT SALT LAKE Michael A. Bowers' Abstract.- The distribution of 21 species of nonvolant mammals among nine islands in the Great Salt Lake was analyzed for biogeographic patterns. The number of species inhabiting an island is closelv correlated with island area That the slope of the regression line describing this relationship (z or b) is relatively shallow compared to (1) totally isolated island systems or (2) island systems ^^'here an equilibrium between rates of colonization and extinc- tion have been attained suggests that isolation plays little role in accounting for the variation in mammalian species diversity among islands. Stepwise multiple regression confirms this, while demonstrating that area alone accounts for 88 percent of the variation in species diversity among islands. However, endemic subs^pecies comprise a significant proportion ot the insular mammalian fauna, suggesting that isolation for small mammals restricted to certain habitats may be substaiitial. A general scenario of the processes determining insular mammalian diversity and endemism is discussed for the Great Salt Lake, where the dynamic lake level creates a potential for different biogeographic pro- cesses over time. o & t- f Analysis of the distribution of nonvolant mammals among oceanic (Carlquist 1965, Wright 1981), landbridge (Hope 1973, South- em 1964, Blocker 1967, Wright 1981), and montane (Brown 1971, 1978) islands has played a significant role in the development and testing of contemporary biogeographic theory (reviews by Brown 1978, Wright 1981). However, most emphasis in the cur- rent biogeographic literature focuses on the distribution of reptiles and birds (Carlquist 1965, Simberloff 1974, V/right 1981). Al- though the conspicuousness of tliese verte- brates facilitate the quantification of species in insular habitats, the poor dispersal abilities of nonvolant mammals (Carlquist 1965, Brown 1971, 1978, Wright 1981) make them particularly interesting subjects for bio- geographic analysis. Isolated habitats in the Intermountain Re- gion of western North America have pro- vided several good tests of biogeographic theory (review by Harper and Reveal 1978). Rigorous interpretation of such patterns have been possible because the paleoclimatic his- tory of the region is well understood (Hubbs and Miller 1948, Martin and Mehringer 1965, Wells and Berger 1967, Wells and Torgensen 1964). This paper discusses the distribution of nonvolant mammals among islands in the Great Salt Lake in relation to the theory of insular biogeography. Special emphasis re- lates distributional patterns to the process of extinction and dispersal, which appear to re- flect historical events rather than equilibrial processes. Methods The distribution of nonvolant mammalian species among nine islands in the Great Salt Lake was compiled from published literature (Stansbury 1852, Fremont 1850, Durrant 1936, 1952, Goldman 1939, Marshall 1940). One source (Marshall 1940), which reported the findings of an extensive mammal survey of the Great Salt Lake in the late 1930s, not only documented the distribution of species (and subspecies) by islands, but also charac- terized in detail the islands proper. Both his- torical (age of island, number of years con- nected to the mainland between 1850 and 1940) and physical (area, island height, dis- tance to mainland) features were included in the general description of each island. Mar- shall (1940) and Durrant (1936, 1952) were particularly interested in the distribution of subspecies restricted to certain islands in the Great Salt Lake. Although both of these au- thors invoked historical explanations to ac- count for the distribution of mammals in gen- eral and endemic subspecies in particular, little emphasis was given to the quantitative 'Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona 85721. 589 590 Great Basin Naturalist Vol. 42, No. 4 analysis of determinants affecting the dis- persion of species and subspecies over the nine islands. In the present paper I reanalyze the baseline data reported by Marshall (1940) in a quantitative manner and interpret the results in light of recent contributions to the general theory of biogeography. The general taxonomy follows Goldman (1939) and Dur- rant (1952). Introduced mammalian species and all chiropterans were excluded from the analysis. For a more detailed account of the islands, see Marshall (1940). Results and Discussion As of 1938, 21 species of non volant mam- mals were distributed among the nine islands (Table 1) of the Great Salt Lake. Moreover, nine subspecies were totally restricted in dis- tribution to these islands. The number of mammalian species (Table 1) inhabiting an island is closely correlated with the area (Table 2) of the island (Fig. 1). When both variables are plotted logarith- mically, the data are well described (r = .74) by a straight line with a slope (z) of .19. Tra- ditionally, biogeographers have expressed this relationship as S = CA^ where S is the num- ber of species inhabiting an island and A is the insular area. The value of the constant (C) and slope (z) have been shown to be de- pendent on the specific taxon and group of islands under consideration (Mac Arthur and Wilson 1967, Preston 1962, Wright 1981). Widespread use of the power function has proved beneficial in that it facilitates the comparison of slopes and, in an a priori sense, biogeographical processes that are impacting insular systems (reviews by Brown 1978, Wright 1981). Table 1. Distribution of subspecies, by island, for mammals in the Great Salt Lake. Taxonomy from Goldman (1939) and Durrant (1952). O Dipodomys microps russeohis' Dipodomys microps alfredi' Dipodomys microps siibtenuis Dipodomys ordii marshalli Dipodomys ordii cineraceus" Dipodomys ordii iitahensis Perognathus parvus plerus" Thomomys bottae minimus" Thomomys bottae nesophilns' Eutamias minimus pictus Spennophilus townsendii mollis Erethizon dorsatum epixanthum Peromyscus crinitus pergracilis Peromyscus maniculatus sonoriensis Peromyscus maniculatus gunnisoni' Peromyscus maniculatus inclarus' Reithrodontamys megalotis ravus Neotoma lepida marshalli' Neotoma lepida lepida Onychornys leucogaster utahensis Sylvilagus nuttalli grangeri Lepus californicus deserticola Odocoileus hemionus hemionus Antiocapra americana americana Lynx rufus pallescens Mustela frenata nevadensis Taxidea taxus taxus Mephitis mephitis inajor Cants latrans lestes Total species X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X X 19 13 'Subspecies not found on the mainland. December 1982 Bowers: Insular Biogeography 591 The z- value (.19) for nonvolant mammals inhabiting islands in the Great Salt Lake is slightly lower than insular biotas where an equilibrium between rates of colonization and extinction have been attained (.20- .35, MacArthur and Wilson 1967). However, the z-value for insular habitats in the present analysis is larger than those commonly re- ported for continental situations (.12-. 17, MacArthur and Wilson 1967) and is com- parable to two decimal places the value re- ported in a recent vegetational analysis of a pinyon-juniper ecosystem where nested quad- rants were used as islands (Harner and Harper 1976). The slight deviation from predicted z- values for continental and equilibria! insular systems can be elucidated through stepwise multiple regression analysis of island charac- teristics (Table 2) on the number of resident mammal species (Table 1). Similar multi- variate techniques have been extremely pow- erful when investigating the significance of interdependent island characteristics (Table 3) on the variation in insular species diversity (Brown 1971, 1978, Harper et al. 1978). That (a) approximately 88 percent of the variation in insular mammalian diversity can be accounted for by island area (Table 4), and (b) there appears to be little impoverishment of species number resulting from isolation by distance to the mainland (Table 4) suggest that colonization rates are high and that there is little effect of insular isolation on species richness. These results appear to be consistent with those of others (Johnson 1975, Brown 1978) studying boreal birds in inter- mountain habitats where avian diversity is at- tributable primarily to the diversity of avail- able habitats and not area per se. Specifically, the overwhelming dominance of area in the multiple regression analysis (Table 4) is in all probability an illusion. Wyckeroff (1973) and Harner and Harper (1976) have demonstrated that both environ- mental favorability and heterogeneity exert a strong influence on the number of vascular plant species per unit area. Because area alone subsumes all these variables, it by itself accounts for several variables that a priori could account for a significant amount of the variation in mammalian species diversity. Unfortunately, data on environmental favor- ability and heterogeneity are not available for the islands in the Great Salt Lake. Quan- tification and analysis of habitat diversity patterns of the islands would be a strong test of the effect of area, per se, on variation in the number of mammalian species in this sys- tem of islands. On the islands of the Great Salt Lake, re- current colonization appears to be much more important than local extinction. This contention can be examined in more detail by looking at ecological attributes of the 21 species that are distributed among the nine islands. In particular, the documentation of characteristics that might increase the proba- bility of local extinction could produce a fin- er degree of resolution than examining entire island compositions. Table 2. Historical and physical data for the nine islands in the Great Salt Lake. Data adapted from Marshall Island Area (hectares^) Dolphin Gunnison Bird (Hat) Sand bar Carrington Badger Stansbury Antelope Fremont 20.7 67.4 9.1 .2 730.2 2.5 7,977..3 10,766.9 1,216.9 Distance' (km) 5.67 24..30 32.40 37.26 32.40 35.64 19.44 5.27 24.30 Height- (m) 18.5 46.2 12.3 .6 123.0 3.1 769.2 737.8 244.9 'Distance (km) from island to the nearest mainland area. 'Maximum height (m) of an island relative to zero ( = 1291.3m) on the Saltair gage. 'Height (m) of connecting bar on the Saltair gage. 'Years an island was connected to the mainland between 1850 and 1938. 'Probable age of an island (see Marshall 1940). Corridor' height (m) Years^ connected .77 -1.23 .31 .31 .31 .31 3.01 .92 .46 18 0 12 7 12 13 76 19 2 Age' (years) 20,000 20,000 20,000 6 20,000 50 70,000 70,000 70,000 592 Great Basin Naturalist Vol. 42, No. 4 o o CO u U u CL CO Li- O cc u CD 3 1.30 1 1.04- 2.0 3.0 AREA- LOG (Hectares^) Fig. 1. The relationship between insular area and the number of nonvolent mammal species for the nine islands in the Great Salt Lake. The equation for the fitted regression and the amount of variance in mammalian species diver- sity accounted for by area (r^) are indicated. The probability of a species becoming ex- tinct appears to be directly dependent on lo- cal population size (MacArthur and Wilson 1963, 1967). In addition to island size, three ecological variables appear to be particularly important in determining insular population size. These variables— body size, trophic sta- tus, and degree of habitat specialization- affect the distribution of mammalian species among the Great Salt Lake islands in the manner we would expect from considerations of their effects on population size: small mammals are foimd on more islands than large ones (Fig. 2), herbivores are better represented than granivores, insectivores, or carnivores, and species that can live only in restricted habitats inhabit fewer islands than species with the same trophic affinities but who are habitat generalists. Figure 2 shows the relationship between the number of is- lands inhabited by a particular species and the logarithm of body weight. Generally these variables do not covary to any signifi- cant extent (r = -.28). However, within the herbivore guild (Fig. 2) the number of islands inhabited by a species is negatively (P<.04) correlated with the logarithm of body weight (y = .64-.09x, r = -.65, df=7). Within this guild, the species of largest body size occur only on the larger islands (Tables 1 and 2), December 1982 Bowers: Insular Biogeography 593 Table 3. Correlation coefficients (r) between variables for the nine islands in the Great Salt Lake. Matrix is com- puted with untransformed data. Note that the number of total species and endemic subspecies are not always closely correlated with the same variables. Corridor Years Number of Area Distance Height height connected Age species Area _ Distance -.593 Height .970 -.547 Corridor height .630 -.279 .677 _ Years connected .610 -.272 .705 .941 _ Age .772 -.583 .859 .402 .466 _ Number of species .938 -.605 .924 .789 .766 .676 Number of endemic subspecies .240 .013 .395 .663 .820 .663 .467 suggesting the importance of matching body size (and, hence, population size) with suit- able habitat patch size, which appears to covary with island area. In fact, those species of both large and small body size that occur on only a few islands usually are found only on large islands (Tables 1 and 2). Thus, it ap- pears that, for mammals on islands in the Great Salt Lake, insular area not only affects the number of species but also can be used to predict some ecological attributes of the denizens. The preceding analyses suggest that isola- tion of insular habitats in the Great Salt Lake is relatively unimportant in determining mammalian diversity. However, the fact that such a high proportion of the island faunas are composed of endemic subspecies strongly suggests that isolation has been a prominent factor, at least in the past, in the evolution of these insular biotas. The determinants pro- moting endemism can be examined by using the proportion of subspecies endemic on each island (Table 1) as the dependent variable and island characteristics as the independent variables (Table 2) in a stepwise multiple regression. The results of this analysis (Table 5) show that two variables (height of the cor- ridor connecting the various islands with the mainland and number of years between 1850 and 1938 that an island was connected to the mainland) explain 76 percent of the variation in the proportion of insular subspecies that are endemic. Obviously, the degree of isola- tion as reflected by corridor height and years connected to the mainland are imminently important in producing and maintaining en- demic subspecies in insular mammalian faunas. It should be noted that in this particu- lar analysis subspecies are considered endem- ic if the corresponding mainland species are of different subspecies. Consequently, endem- ic subspecies can and do occur on several islands. In the analysis of species distributed among islands, isolation played little role in accoimt- ing for the patterns. In contrast, the occur- rence of endemic subspecies is highly corre- lated with the degree of isolation. These differences can be reconciled by examining ecological attributes of the endemic sub- species. That all endemic subspecies are her- bivores, granivores, or omnivores and are rel- atively small (<300g) when compared with the other species that are distributed among the nine islands suggests that population size may effect the rate of local genotypic differ- entiation from mainland populations. Specifi- cally, the data suggest that species capable of maintaining large insular populations per Table 4. Summary of stepwise multiple regression of the influence of island characteristics on the number of mammalian species on islands in the Great Salt Lake. Variable" Order entered in equation Contribution toR2 Area Corridor height Distance 0.882 0.065 0.008 "Data from Table 2 (untransformed). F-value 52.16 7.36 .96 Significance level .0001 .035 .373 594 Great Basin Naturalist Vol. 42, No. 4 8 t 7 Q LxJ CD < I Z LO Q Z < 6 - 5 - t/) 4 - O cc UJ CD 3 3 - 2 - PM o DO a y= 5.2 ♦(- .70)X r2 = .08 P> .05 CL pp TB MM EC ^"^ .,„^^_^ OM a • £^ • ^^"^ • RM PCOL CM MF TT LR AA O o ■ o C^ £^ ^ • • 1 r "■ 1 T ■ 1 1 1 10 1.6 2.2 23 3.4 4.0 4.6 52 BODY WEIGHT - LOG (grams) Fig. 2. Frequency of occurrence of species of nonvolent mammals on the nine islands in the Great Salt Lake plotted against body weight; solid circles represent herbivores, hollow circles represent omnivores, hollow squares represent granivores, hollow triangles represent carnivores, and solid squares represent insectivores. Letters adjacent to each point refer to species names that can be identified by reference to Table I. unit area are more likely to show substantial differentiation from the mainland popu- lations than species with smaller populations. Two interdependent factors may account for these differences. First, as a consequence of larger body size, and thus mobility, more colonists of large-bodied species probably dis- perse to islands than smaller species. Thus, the amount of gene flow from mainland to is- land populations would be relative to body size: e.g., because of intrinsic differences in dispersal ability insular populations of large species would be expected to have gene fre- quencies closer to those mainland popu- lations when compared to smaller species. Second, given that a single colonist did dis- perse to an island from the mainland, the in- corporation and ultimate fixation of its genes into the genepool of the population would be a negative function of population size. Spe- cifically, the larger the population the great- er the likelihood that a single colonist's con- tribution of novel genes will be swamped: e.g., the infusion of new genetic material from one act of colonization will be much greater in a population of 10 individuals when compared to a population of 100. Although the analyses presented in this present paper generally suggest that isolation has not been important in limiting the distri- bution of nonvolant mammals among the is- lands in the Great Salt Lake, a caveat should be interjected. Specifically, the survey by Marshall (1940) was conducted when the lake December 1982 Bowers: Insular Biogeography 595 Table 5. Summary of stepwise multiple regression of the influence of island characteristics on the proportion of subspecies that are endemic for the nine islands in the Great Salt Lake. Variable" Order entered in equation' Corridor height Years connected Age "Data from Table 2 (untransformed). Contribution toR2 .16 .60 .07 F-value 1.32 14.64 2.08 Significance level .287 .009 .208 level was near a 100-year minimum. Coloni- zations by mammals just prior to the survey could effectively obscure patterns of local ex- tinctions, thereby downplaying the role of isolation. Since that time, the lake level has remained relatively high. A strong test of the result presented here would be to re-census the islands while test- ing for the effects of isolation. Not only would this yield valuable biogeographical in- formation, but it would serve as one of the few instances where island relaxation rates (Diamond 1972, 1975) could be precisely quantified because the exact year of isolation by island is known through the detailed docu- mentation of the lake level (U.S. Weather Bureau). General Discussion The distribution of nonvolant mammals among nine islands in the Great Salt Lake ap- pears to reflect the effect of recurrent coloni- zations rather than the equilibrial processes of extinction and immigration. Consequently, islands appear to be "saturated" with species and the distribution of species among islands is probably determined by the distribution of amicable habitats that are extensive enough to support populations of the colonists. How- ever, isolation of insular habitats does appear to be important for some species of small body size. This is supported by a high degree of endemism for these species. The failure of the equilibrium theory of biogeography (MacArthur and Wilson 1963, 1967) to account for the distribution of mam- mals among insular habitats studied here sup- ports a general trend that contrasts inter- mountain insular habitats with oceanic island systems. Specifically, the distribution of birds (Johnson 1975, Behle 1978), mammals (Brown 1971, 1978), fish (Smith 1978) and even plants (Harper et al. 1978) among insular habitats in western North America rarely conform to equilibrial predictions. In con- trast, the biotas of oceanic islands often ex- hibit equilibrial distributions (Simberloff 1974, Wright 1981). This difference is corre- lated with contrasting paleoclimatic and geo- logical processes impacting the two types of insular systems. Over the last million years extensive climatic and geological changes have drastically changed and, for islands in the Great Salt Lake, are still changing the landscape of western North America (see ear- lier citations of Hubbs, Martin, Wells). Oce- anic systems, however, have a long history of isolation and relative environmental stability (MacArthur and Wilson 1967, Simberloff 1974). It is possible that these differences in the underlying environmental patterns (both past and present) account for the contrasting biogeographic tendencies. As pointed out by Brown (1978), these dynamic environmental factors should differentially affect species with contrasting vagilities. Because mammals in general are relatively poor dispersers across unsuitable habitats (Brown 1971, 1978) and aquatic barriers (Carlquist 1965), it is in- teresting to speculate what biogeographic model the Great Salt Lake islands would con- form to if the islands were isolated for long periods of time. In all probability the mainte- nance of a high lake level would create a sys- tem analogous to that of boreal mammals on mountain tops (Brown 1971, 1978), where ex- tinctions are dependent on insular size and colonizations are rare. Such processes may currently be in effect with the higher lake level. Acknowledgments Initial interest in the quantification of mammals distributed among islands in the Great Salt Lake was generated through dis- cussions with T. C. Gibson and J. H. Brown. 596 Great Basin Naturalist Vol. 42, No. 4 The present analyses would have been impos- sible without the pioneering work of natural historians and systematists who documented in detail the mammalian fauna of the Great Salt Lake area. To all the above I am grateful. Literature Cited Behle, W. H. 1978. Avian biogeography of the Great Basin and Intermountain Region. Great Basin Nat. Mem. No. 2:55-88. Bloeker, J. C. VON, Jr. 1967. Land mammals of the southern CaHfornia islands. R. N. Philbrick, ed.. Proceedings of the Symposium on the Biology of the California Channel Islands. Santa Barbara Botanic Garden. Santa Barbara, California, USA. Brown, J. H. 1971. Mammals on mountaintops: non- equilibrium insular biogeography. American Nat- uralist 105:467-478. 1978. The theory of insular biogeography and the distribution of boreal birds and mammals. Great Basin Nat. Mem. No. 2:209-227. Carlquist, S. 1965. Island life. Natural History, New York, New York. Diamond, J. M. 1972. Biogeographic kinetics: estimation of relaxation times for avifaunas of southwest Pacific islands. Proc. U.S. Nat. Acad. Sci. 69:3199-320.3. 1975. Assembly of species communities. Pages 342-444 in M. L. Cody and J. M. Diamond, eds.. Ecology and evolution of communities. Harvard University Press, Cambridge, Massachusetts, USA and London, England. DuRRANT, S. D. 1936. A new gopher from Antelope Island, Great Salt Lake, Utah. J. Mammal. 20:351-357. 1952. Mammals of Utah: taxonomy and distribu- tion. Univ. Kansas Publ. Lawrence, Kansas, USA. Fremont, J. C. 1850. The exploring expedition into the Rocky Mountains, Oregon California. Derby and Co., Buffalo, New York, USA. Goldman, E. A. 1939. Nine new mammals from islands in the Great Salt Lake, Utah. J. Mammal. 20:351-357. Harner, R. F., and K. T. Harper. 1976. The role of area, heterogeneity, and favorability in plant spe- cies diversity of pinyon-junior ecosystems. Ecolo- gy 57:1254-1263. 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. No. 2:81-103. Harper, K. T., and J. L. Reveal, eds. 1978. Inter- mountain biogeography: a symposium. Great Ba- sin Nat. Mem. No. 2:1-268. Hope, J. H. 1973. Mammals of the Bass Straight Islands. Proc. Roy. Soc. Victoria 85:163-195. HuBBS, C. L., AND R. R. Miller. 1948. Correlation be- tween fish distribution and hydrographic history in the desert basins of western United States. Pages 17-166 in The Great Basin, with emphasis on glacial and postglacial times. Bulletin of the Univ. of Utah 38:1-191. Johnson, N. K. 1975. Controls of numbers of bird spe- cies on montane islands in the Great Basin. Evo- lution 29:545-557. MacArthur, R. H., and E. O. Wilson. 1963. An equi- librium theory of insular zoogeography. Evolu- tion 17:373-387. 1967. The theory of island biogeography. Prince- ton Univ. Press, Princeton, New Jersey. Marshall, W. H. 1940. A survey of the mammals of the islands in Great Salt Lake, LItah. J. Mammal. 21:144-159. Martin, P. S., and P. J. Mehringer. 1965. Pleistocene pollen analysis and biogeography of the South- west. Pages 435-451 in H. E. Wright and D. G. Frey, eds.. The quarternary of the United States. Princeton LIniv. Press, Princeton, New Jersey, USA. Preston, F. W. 1962. The canonical distribution of com- monness and rarity: Part I. Ecology 43:185-215. SiMBERLOFF, D. S. 1974. Equilibrium theory of island biogeography and ecology. Ann. Rev. Ecol. Syst. 5:161-182. Smith, G. R. 1978. Biogeography of intermountain fishes. Great Basin Nat. Mem. No. 2:17-42. Southern, H. N. 1964. The handbook of British mam- mals. Blackwell Scientific, Oxford, England. Stansbury, H. 1852. Exploration and survey of the Val- ley of the Great Salt Lake of Utah. Lippincott, Gromlee and Co., Philadelphia, Pennsylvania, USA. U.S. Weather Bureau. 1979. Cliniatological data (series by states). Wells, P. V., and R. Berger. 1967. Late Pleistocene history of coniferous woodland in the Mojave Desert. Science 155:1640-1647. Wells, P. V., and C. D. Jorgensen. 1964. Pleistocene woodrat middens and climatic change in the Mo- jave Desert: a record of juniper woodlands. Sci- ence 143:1171-1173. Wright, S. J. 1981. Intra-archipelago vertebrate distri- butions: the slope of the species-area relation. American Nat. 118:726-748. Wyckoff, J. W. 1973. The effect of soil texture on spe- cies diversity in an arid grassland of the eastern Great Basin. Great Basin Nat. 33:163-168. DORSAL HAIR LENGTH AND COAT COLOR IN ABERT'S SQUIRREL {SCIURUS ABERTT) Denis C. Hancock, Jr.', and Donald J. Nash' Abstract.- Sciurus ahertl like many other sciurids including Sciurus vulgaris and Sciurus carolinensis shows a coat color polymorphism. Like Sciurus vulgaris, Sciurus aberti shows a correlation between coat color phase and dor- sal hair length. Both squirrels show an increased frequency of dark morphs in the northern portions of their respec- tive ranges. ^ A number of sciurids, including Sciurus aberti (Ramey and Nash 1976), Sciurus caroli- nensis (Creed and Sharp 1958), and Sciurus vulgaris (Voipio 1956) show coat color poly- morphisms. The genetics of these polymor- phisms seem to involve mutations at the ex- tension locus as with Sciurus vulgaris, mutations at the agouti locus as with Sciurus carolensis, or mutations at both loci as with Sciurus aberti (Searle 1968, Ramey and Nash 1976). For the above three species, field ob- servation and sampling have established that the darker morphs are more common in the northern reaches of their respective ranges or at higher elevations. Voipio and Hissa (1970) measured hair density and length and related them to pel- age color. They found significant differences for hair densities, hair lengths, and hair weights and concluded that hair density was greater for the dark phase of the European Red Squirrel (Sciurus vulgaris) than for the light (red) phase. The present study was designed to examine the relationship between hair length and coat color in Abert's Squirrel (Sciurus aberti fer- reus). This subspecies occurs in north central Colorado and is the best documented for col- or polymorphism. It occurs in two main phases: gray and nearly black as well as some intermediate phenotypes (Nash and Seaman 1977). The gray morphs typically show an agouti pigment distribution and the black forms are typically nonagouti. If the squirrels are classified on the basis of pigment distribu- tion, a color range for each group is demon- strated, with those in the nonagouti group tending to be much darker than those in the agouti group. Dorsal guard hairs and underfurs were measured on 23 specimens of Sciurus aberti ferreus. Because Abert's Squirrels were fully protected in Colorado at the time measure- ments were taken, museum specimens from Colorado State University and the Denver Museum of Natural History were used. A small tuft of hairs was removed at skin level from the middorsal region of each squirrel. Guard hairs were measured three times to the nearest millimeter and averaged, and un- derfurs were measured as a group to the nearest millimeter. Statistical analyses in- cluded mean, standard deviation, and a t-test on the two respective means (agouti and non- agouti squirrels), with the null hypothesis that the two means were the same. Nonagouti Abert's Squirrel guard hairs (n= 14) averaged 23.52 mm in length; Agouti guard hairs (n = 9) averaged 20.92 mm. For underhairs, nonagouti (n= 14) averaged 15.14 mm and agouti (n = 9) averaged 13.00 mm. A T-test was performed on these data and the null hypothesis that the two means were identical was rejected (p<.001). Dark and light pelage appear to be equally effective in heat conservation at lower tem- peratures (Creed and Sharp 1958), so if re- sponse to cold alone were responsible for the maintenance of the polymorphisms discussed here, one could just as likely find a race of lighter than normal squirrels at higher eleva- tions or in northern latitudes, assuming these 'Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado 80523. 597 598 Great Basin Naturalist Vol. 42, No. 4 genes were not deleterious in other ways. However, if genes for longer pelage were as- sociated through linkage or pleiotropism with genes for darker color, the gene or genes could be selected for in colder areas. To confirm this hypothesis, it is necessary to determine the genetic basis for coat color polymorphism more accurately and, having done that, sample the squirrels in their natu- ral habitat while obtaining climatological data for the area of capture. This would de- termine whether or not there is a significant difference in average climate. If there is a link between color phase and hair length in sciurids, it may explain why there do not appear to be many races of lighter than normal squirrels in the northern portions of their ranges. Literature Cited Creed, W. A., and W. M. Sharp. 1958. Melanistic gray squirrels in Cameron County, Pennsylvania. J. Mammal. 39:532-537. Nash, D. J., and R. N. Seaman. 1977. Sciurus aberti. Mammalian Species 80:1-5. Ramey, C. a., and D. J. Nash. 1976. Coat color poly- morphism of Abert's squirrel, Sciurus aberti, in Colorado. Southwest Nat. 21:209-217. Searle, a. G. 1968. Comparative genetics of coat colour in mammals. Academic Press, New York. 308 pp. VoiPio, P. 1956. The biological zonation of Finland as reflected in zootaxonomy. Ann. Zool. Soc. Van- amo 18:1-36. VoiPio, P., and R. Hissa. 1970. Correlation with fur density of color polymorphism in Sciurus vul- garis. J. Mammal. 51:185-187. THE RACCOON, PROCYON LOTOR, IN WYOMING E. Blake Hart' Abstract.- Recent distribution maps show raccoon as occupying only the extreme eastern-northeastern portion of Wyoming. However, there is substantial evidence that raccoon are common throughout Wyoming and currently inhabit all the major and many of the minor drainages throughout Wyoming. The distribution of the raccoon, Procyon lotor, has changed rather dramatically in the past few years in Wyoming. Long (1965) re- corded that it was present in only five east- em and northeastern counties of Wyoming in 1965. In more recent years, however, evi- dence has accumulated suggesting that rac- coon range is on the increase in the state. Hoffmann, Wright, and Newby (1969) de- scribed several localities in adjoining Mon- tana from which raccoon had been taken. Al- though Hall's (1981) summary showed nearby records in adjacent areas of Idaho, Utah, and Colorado, he also indicated no raccoon local- ities in central and western Wyoming. Clark, Saab, and Casey (1980) in their review of Wyoming mammal literature likewise pre- sented no new citations on current raccoon distribution. Lotze and Anderson (1979) pre- sented no range extensions. Dale Weston, a rancher in rural Sage (20 miles south, 2 miles west, Cokeville), Lincoln Co., trapped a raccoon during the winter of 1980-1981. He communicated instances of road kills and had knowledge of trappers who had taken raccoon locally. Upon further in- quiry, including written and verbal commu- nication with several fur dealers in western and central Wyoming, we found that the rac- coon is presently a common mammal throughout the state and has been for some time, especially the past 5 to 15 years. Matt Failoni, a fur buyer from Kemmerer, Lincoln Co., stated that he had purchased 51 raccoon pelts around the Kemmerer area in the past five years. James Cook, a fur dealer in Evanston, Uintah Co., had purchased ap- proximately 20 raccoon each year for the past five years. Recently, Charles Neely, a fur buyer and trapper from Pinedale, Sublette Co., person- ally trapped 12 raccoon. Due to the paucity of trappers in that area, fewer raccoon have been taken than might otherwise be ex- pected. Beaver trappers occasionally have taken raccoon, but, due to the reduced prices of pelts, fewer beaver traps have been set. Mr. Neely recounted that although few local Pinedale residents raise chickens, every one has called him and complained about rac- coon harassment. Mr. Neely further said that his uncle trapped a raccoon locally 30-35 years ago, but in only the past five or so years have numbers of animals increased appre- ciably. He estimated that extant distribution of raccoon averages one for every three to four miles of creek and river bottom, a total of 30 to 40 raccoons within a 20 mile radius of Pinedale. Mr. Neely suggested that rac- coons might have immigrated into the Pine- dale area from the south up the Green River drainage system. Jake Korell, Riverton, Fremont Co., pur- chased approximately 300 raccoon pelts dur- ing the 1981-1982 season and an average of 450 for each of the previous five years, or a total of about 2000 animals. Mr. Korell stated that 46 years ago there were no raccoon in the Riverton area, that they first appeared about 30 years ago. He suggested that con- temporary local populations originated both from accidental liberation of pet raccoons by a Missouri family and also from immigration of wild raccoon up local drainage systems. 'Department of Mathematics and Natural Sciences, Northern State College, Aberdeen, South Dakota 57401. 599 600 Great Basin Naturalist Vol. 42, No. 4 Mr. Korell also said that the severe winter of 1978-1979 caused significant mortality, that present numbers were down somewhat. Herman Genz, Rawlins, Carbon Co., trapped and purchased about eight raccoon during the past five years. He caught none in the 1981-1982 fur season, but trapped two or three per year in previous years in the Pass Creek area; he also purchased seven or eight local raccoon in the past five years. Ronald Yates, a statewide fur buyer from Casper, Natrona Co., purchased 300 raccoon pelts in the 1981-1982 season and approx- imately 1500 locally during the past five years. He also purchased about 50 in 1981-1982, 300 to 400 in the past five years, out of the Green River drainage system. He stated that raccoons are now being taken where none have been found previously. Mr. Yates stated unequivocally that raccoon are currently present in all of the major drainage systems in the state of Wyoming, bar none. Raccoon from the Snake River drainage are of especially high quality, and there seems to be an abundance of animals along the Big Horn River in the general vicinity of Thermopolis, Hot Springs Co. He also sug- gested that the Shoshone River may well have served as a dispersal corridor into Wyoming from the Big Horn River, Mon- tana, where, as a boy, he was aware of abun- dant raccoon populations. In summary, there is strong evidence that significant populations of raccoon presently occur in areas of suitable habitat throughout the state of Wyoming. Literature Cited Clark, T. W., V. A. Saab, and D. Casey. 1980. A par- tial bibliography of Wyoming mammals. North- west Sci. 54(l):55-67. Hall, E. R. 1981. Mammals of North America. John Wiley and Sons, N.Y. 1181 pp. Hoffmann, R. S., P. L. Wright, and F. E. Newby. 1969. The distribution of some mammals in Mon- tana. J. Mammal. 50(3):579-604. Long, C. A. 1965. The mammals of Wyoming. Univ. Kansas. Publ., Mus. of Nat. Hist. 14(18):493-758. LoTZE, J. H., AND S. Anderson. 1979. Procijon lotor. Mamm. Species 119:1-8. INTERCANINE CROWN DISTANCES IN RED FOXES AND BADGERS E. Blake Hart' Abstract.- Intercanine crown distances of 605 wild South Dakota red foxes and 249 badgers of unknown age were measured; adults and juveniles were diagnosed by radiographs of canine teeth. In foxes, T-tests between similar age, between similar sex, and between combined age groups were significant at .01. In badgers, significance was found only between adult males and juvenile males and between adult males and adult females at .05. With refinement of fiirbearer management practices, methodology of accurate age de- termination is paramount in obtaining infor- mation on current population status. Various workers (Churcher 1960, Grue and Jensen 1976, MacPherson 1969, Morris 1972) have shown that a variety of cranial characters are correlative with the aging process in carnivores. In tile red fox, for example, several skull characters have been shown by some of the above to be reliable indicators of age, such as closure of various sutures, triangularity of postorbital processes, texture of temporal areas, pointedness of nasals, and several den- tal characters that include numbers of in- cremental annuli, pulp cavity size, enamel line distance, overall tooth wear, etc. Churcher (1960) was able to differentiate with a fair degree of accuracy between sexes of similarly aged fox by graphing mastoid width against total skull length X zygomatic width. An additional character that was thought to possibly have correlation with aging in wild populations of red foxes and badgers is the distance between crowns of normally rooted canine teeth. Upper jaws of over 600 unknown age red foxes and lower jaws of 249 unknown age badgers were obtained from a local fur dealer and were cut from skulls with pruning shears. Measurements of maxillary intercanine crown distances were then made with vernier calipers to the nearest 0.1 mm. Jaws were boiled to loosen canines, after which teeth were removed and X-rayed to distinguish be- tween juveniles and adults. Results were then analyzed to determine significance (Figure !)• In foxes, all T-tests computed between sim- ilar age, similar sex groups, and combined age groups were significant at 0.1; a T-test between adult females and juvenile males was not significant at .05. In badgers, T-tests between adult and juve- nile males and adult males and adult females were significant at .05. No significance (.05) was encountered between adult females and juvenile females, between juvenile males and juvenile females, and between combined adults and combined juveniles. This study was based upon unknown age South Dakota carnivores that were parti- tioned as either adults or juveniles by relative size (X-ray) of pulp cavity, an accepted tech- nique among many wildlife workers. These results should be interpreted with the knowl- edge that relative pulp cavity size has yet to be shown as absolute. It is hoped that these results will stimulate further studies of intercanine crown distances in known age wild foxes and badgers. Literature Cited Churcher, C. S. 1960. Cranial variation in the North American red fox. J. Mammal. 41(3):349-360. Grue, H., and B. Jensen. 1976. Annual cementum struc- tures in canine teeth in arctic foxes (Alopex la- gopsusil). Danish Rev. of Game Biol. 10(3):1-12. Morris, P. 1972. A review of mammalian age determi- nation. Mamm. Rev., 2:69-104. MacPherson, A. H. 1969. The dynamics of Canadian arctic fox populations. Canadian Wildl. Serv. Rept., Series 8. 52 pp. 'Department of Mathematics and Natural Sciences, Northern State College, Aberdeen, South Dakota 57401. 601 602 Great Basin Naturalist Vol. 42, No. 4 28 27 S 26 Q 25 Z o tt 24 u LU z Z 23 < u 22 21 a^5~m fa^~f ijuv m ijuv f tall ad lall juv FOX AGE & SEX GROUPS Fig. 1. Maxillary intercanine crown distances of wild South Dakota red foxes. Adults were partitioned from juve- niles by X-rays of canine teeth. Means, longer horizontal lines; two standard errors, boxes; and two standard devia- tions, vertical lines. INDEX TO VOLUME 42 The genera and species described as new to science in this volume appear in bold type ii this index. A description of Timpie Springs, Utah, with a preliminary survey of the aquatic macrobiota, p. 77. A new species of Cryptantha (Boraginaceae) from Nevada, p. 196. A new species of Penstemon (Scrophulariaceae) from the Uinta Basin of Utah and Colorado, p. 367. A species of Cryptantha (Boraginaceae) dedicated to the memory of F. Creutzfeldt, p. 203. Additions to the vascular flora of Montana and Wyoming, p. 413. Algal populations in Bottle Hollow Reservoir, Duchesne County, Utah, p. 205. Allred, Dorald M., article by, p. 415. Allred, Kelly Wayne, article by, p. 101. Amphicraniis splendens, p. 223. Andersen, Ferron L., Lauritz A. Jensen, and Jordan C. Pederson, article by, p. 351. Andersen, Ferron L., Lauritz A. Jensen, and Peter M. Schantz, article by, p. 65. Ants of Utah, p. 415. Araptus micropilosus, p. 224. Araptus morigerus, p. 224. Araptus placetulus, p. 225. Austin, D. D., and Philip J. Umess, article by, p. 512. Austin, George T., and J. Scott Miller, article by, p. 232. Avery, David F., and Wilmer W. Tanner, article by, p. 273. Baugh, Thomas M., Michael A. Nelson, and Floyd Simpson, article by, p. 77. Behavior and habitat preferences of ring- necked pheasants during late winter in central Utah, p. 562. Bowers, Janice E., article by, p. 105. Bowers, Michael A., article by, p. 589. Briggs, George M., and James A. MacMahon, article by, p. 50. Brotherson, Jack D., article by, p. 246. Brotherson, Jack D., and Jeffrey G. Skousen, article by, p. 562. Brotherson, Jack D., and Miles O. Moretti, article by, p. 81. Brotherson, Jack D., Joy D. Cedarleaf, and S. Dick Worthen, article by, p. 91. Buccal floor of reptiles, a summary, p. 273. Campbell, Thomas M., Ill, Tim W. Clark, and Craig R. Groves, article by, p. 100. Campbell, Thomas M., Ill, Tim W. Clark, David G. Socha, and Denise E. Casey, article by, p. 572. Campos, E. G., and R. B. Eads, article by, p. 241. Carter-Lovejoy, Steven H., article by, p. 113. Casey, Denise E., Tim W. Clark, Thomas M. Campbell III, and David G. Socha, article by, p. 572. Cedarleaf, Joy D., S. Dick Worthen, and Jack D. Brotherson, article by, p. 91. Chatterley, L. Matthew, Blaine T. Welsh, and Stanley L. Welsh, article by, p. 385. Chramesus bispinus, p. 225. Cirsium eatonii var. harrisonii, p. 200. Cirsium eatonii var. murdockii, p. 200. Cirsium ownbeyi, p. 200. Cirsium scariosum var. thorneae, p. 201. Cirsium virginensis, p. 201. Clark, Tim W., Thomas M. Campbell III, and Craig R. Groves, article by, p. 100. Clark, Tim W., Thomas M. Campbell III, David G. Socha, and Denise E. Casey, article by, p. 572. Cnesinus aquihuai, p. 226. Cnesinus maris, p. 226. Cooper, James J., article by, p. 60. Courtenay, Walter R., Jr., and James E. Deacon, article by, p. 361. Crataegus douglasii var. duchesnensis, p. 10. Creasy, James, and Robert B. Finley, Jr., article by, p. 360. Cryptantha creutzfeldtii, p. 203. 603 604 Great Basin Naturalist Vol. 42, No. 4 Cryptantlxa welshii, p. 196. Deacon, James E., and Paul Gregor, article by, p. 549. Deacon, James E., and Walter R. Courtenay, Jr., article by, p. 361. Dendrocranulus gracilis, p. 227. Description of a new Phalacropsylla and notes on P. alios (Siphonaptera: Hystrichopsyllidae), p. 241. Description of the female of Phalacropsylla hamata (Siphonaptera: Hystrichopsyllidae, p. 96. Diameter-weight relationships for juniper from wet and dry sites, p. 73. Distribution and relative abundance of fish in Ruth Reservoir, California, in relation to environmental variables, p. 529. Distribution of the moss family Grimmiaceae in Nevada, p. 583. Dom, Robert D., and Robert W. Lichvar, article by, p. 413. Dorsal hair length and coat color in Abert's squirrel {Sciurus aberti), p. 597. Eads, R. B., and E. G. Campos, article by, p. 241. Eads, R. B., and G. O. Maupin, article by, p. 96. Early development of the razorback sucker, Xyrauchen texanus (Abbott), p. 553. Ecomorphology and habitat utilization of Echinocereus engelmannii and E. triglochidiatus (Cactaceae) in southeastern California, p. 353. Effects of defoliation on reproduction of a toxic range plant, Zigadenus paniculatus, p. 524. Emmerson, Frederick H., article by, p. 350. England, John Larry, article by, p. 367. Finley, Robert B., Jr., and James Creasy, article by, p. 360. First record of pygmy rabbits {Brachylagus idahoensis) in Wyoming, p. 100. First specimen of the spotted bat {Eudemia maculatum) from Colorado, p. 360. Flinders, Jerran T., William D. Steigers, Jr., and Susan M. White, article by, p. 567. Greger, Paul, and James E. Deacon, article by, p. 549. Groves, Craig R., Thomas M. Campbell III, and Tim W. Clark, article by, p. 100. Growth of juvenile American lobsters in semiopen and closed culture systems using formulated diets, p. 67. Gustafson, Eric S., and W. L. Minckley, article by, p. 553. Habitat manipulation for reestablishment of Utah prairie dogs in Capitol Reef National Park, p. 517. Hancock, Denis C, Jr., and Donald J. Nash, article by, p. 597. Hansen, Richard M., and James G. MacCracken, article by, p. 45. Hart, E. Blake, articles by, p. 599, 601. Hassler, Thomas J., and Steven Vigg, article by, p. 529. Heckmann, R. A., S. R. Wadley, R. C. Infanger, and R. W. Mickelsen, article by, p. 67. Herpetological notes from the Nevada Test Site, p. 219. Higgins, Larry C, and Kaye H. Thome, article by, p. 196. Hylastes retifer, p. 227. Hylociinis prolatus, p. 228. Infanger, R. C, S. R. Wadley, R. A. Heckmann, and R. W. Mickelsen, article by, p. 67. Insular biogeography of mammals in the Great Salt Lake, p. 589. Intercanine crown distances in red foxes and badgers, p. 601. Invertebrate faunas and zoogeographic significance of lava tube caves of Arizona and New Mexico, p. 405. Jensen, Lauritz A., Ferron L. Andersen, and Peter M. Schantz, article by, p. 65. Jensen, Lauritz A., Jordan C. Pederson, and Ferron L. Andersen, article by, p. 351. Johansen, Jeffrey, Samuel R. Rushforth, and Irena Kaczmarska, article by, p. 205. Kaczmarska, Irena, Jeffrey Johansen, and Samuel R. Rushforth, article by, p. 205. Krebill, Richard G., and David L. Nelson, article by, p. 262. Lavin, Matt, article by, p. 583. Lichvar, Robert W., and Robert D. Dom, article by, p. 413. Local floras of the Southwest, 1920-1980: an annotated bibliography, p. 105. Lund, R., and T. Weaver, article by, p. 73. MacCracken, James G., and Richard M. Hansen, article by, p. 45. MacMahon, James A., and George M. Briggs, article by, p. 50. Mathiasen, Robert L., article by, p. 120. December 1982 Index 605 Maupin, G. O., and R. B. Eads, article by, p. 96. Mickelsen, R. W., S. R. Wadley, R. A. Heckmann, and R. C. Infanger, article by, p. 67. Micracis burgosi, p. 228. Miller, J. Scott, and George T. Austin, article by, p. 232. Minckley, W. L., and Eric S. Gustafson, article by, p. 553. Moretti, Miles O., and Jack D. Brotherson, article by, p. 81. Murphy, Joseph R., and Dwight G. Smith, article by, p. 395. Nash, Donald J., and Denis C. Hancock, Jr., article by, p. 597. Nelson, David L., article by, p. 369. Nelson, David L., and Richard G. Krebill, article by, p. 262. Nelson, Michael A., Thomas M. Baugh, and Floyd Simpson, article by, p. 77. Nest site selection in raptor communities of the eastern Great Basin desert, p. 395. New species of American bark beetles (Coleoptera: Scolytidae), p. 223. New taxa of thistles {Cirsium: Asteraceae) in Utah, p. 199. Observations on the reproduction and embryology of the Lahontan tui chub, Gih bicolor, in Walker Lake, Nevada, p. 60. Observations on woundfin spawning and growth in an outdoor experimental stream, p. 549. Occurrence and effect of Chrysomyxa pirolata cone rest on Picea pungens in Utah, p. 262. Paspalum distichum L. var. induturn Shinners (Poaceae), p. 101. Peck, Stewart B., article by, p. 405. Pederson, Jordan C., Lauritz A. Jensen, and Ferron L. Andersen, article by, p. 351. Pensternon albifluvis, p. 367. Phalacropsis dispar (Coleoptera: Phalacridae), an element in the natural control of native pine stem rust fungi in the western United States, p. 369. Phloeotrihus perniciosus, p. 229. Player, Rodney L., and Philip J. Umess, article by, p. 517. Prairie dog colony attributes and associated vertebrate species, p. 572. Preliminary index of authors of Utah plant names, p. 385. Prevalence of Elaeophora schneideri and Onchocerca cervipedis in mule deer from central Utah, p. 351. Pseiidothysanoes fimbriatus, p. 229. Pseiidothysanoes pini, p. 229. Rhythm of fecal production and protein content for black-tailed jackrabbits, p. 567. Rushforth, Samuel R., Jeffrey Johansen, and Irena Kaczmarska, article by, p. 205. Schantz, Peter M., Lauritz A. Jensen, and Ferron L. Andersen, article by, p. 65. Scohjtodes pilifer, p. 230. Scolytus binodus, p. 230. Seasonal foods of coyotes in southeastern Idaho: a multivariate analysis, p. 45. Simpson, Floyd, Thomas M. Baugh, and Michael A. Nelson, article by, p. 77. Skousen, Jeffrey G., and Jack D. Brotherson, article by, p. 562. Smith, Dwight G., and Joseph R. Murphy, article by, p. 395. Socha, David G., Tim W. Clark, Thomas M. Campbell III, and Denise E. Casey, article by, p. 572. Species-habitat relationships in an Oregon cold desert lizard community, p. 380. Status of introduced fishes in certain spring systems in southern Nevada, p. 361. Steigers, William D., Jr., Jerran T. Flinders, and Susan M. White, article by, p. 567. Structure of Alpine plant communities near King's Peak, Uinta Mountains, Utah, p. 50. Tanner, Wilmer W., article by, p. 219. Tanner, Wilmer W., and David F. Avery, article by, p. 273. Taxonomic studies of dwarf mistletoes {Arceuthobium spp.) parasitizing Pinus strobiformis, p. 120. Temperature and salinity relationships of the Nevadan relict dace, p. 541. Temperature-related behavior of some migrant birds in the desert, p. 232. Tepedino, V. J., article by, p. 524. The raccoon, Procyon lotor, in Wyoming, p. 599. The prevalence of Echinococcus granulosus and other taeniid cestodes in sheep dogs of central Utah, p. 65. 606 Great Basin Naturalist Vol. 42, No. 4 The relation between species numbers and island characteristics for habitat islands in a volcanic landscape, p. 113. Thome, Kaye H., and Larry C. Higgins, article by, p. 196. Umess, Philip J., and D. D. Austin, article by, p. 512. Umess, Philip J., and Rodney L. Player, article by, p. 517. Utah flora: Rosaceae, p. 1. Utah plant types— historical perspective 1840 to 1981— annotated list, and bibliography, p. 129. Vegetal responses and big game values after thinning regenerating lodgepole pine, p. 512. Vegetation and soil factors in relation to soil position on foothill knolls in the Uinta Basin of Utah, p. 81. Vegetation of the mima mounds of Kalsow Prairie, Iowa, p. 246. Vigg, Steven, article by, p. 541. Vigg, Steven, and Thomas J. Hassler, article by, p. 529. Wadley, S. R., R. A. Heckmann, R. C. Infanger, and R. W. Mickelsen, article by, p. 67. Weather conditions in early summer and their effects on September blue grouse (Dendragapus ohscurus) harvest, p. 91. Weaver, T., and R. Lund, article by, p. 73. Welsh, Blaine T., L. Matthew Chatterley, and Stanley L. Welsh, article by, p. 385. Welsh, Stanley L., articles by, p. 1, 129, 199, 203. Welsh, Stanley L., L. Matthew Chatterley, and Blaine T. Welsh, article by, p. 385. Werschkul, David F., article by, p. 380. Western diamondback rattlesnake in southern Nevada: a correction and comments, p. 350. White, Susan M., William D. Steigers, Jr., and Jerran T. Flinders, article by, p. 567. Wood, Stephen L., article by, p. 223. Worthen, S. Dick, Joy D. Cedarleaf, and Jack D. Brotherson, article by, p. 91. Yeaton, Richard I., article by, p. 353. 7070 029 NOTICE TO CONTRIBUTORS Manuscripts intended for publication in the Great Basin Naturalist or Great Basin Natural- ist Mmioirs 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 margms 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. 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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 repnnt 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 Mermountain 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 Ants of Utah. Dorald M. Allred 415 Vegetal responses and big game values after thinning regenerating lodgepole pine. D. D. Austin and Philip J. Urness 512 Habitat manipulation for reestablishment of Utah prairie dogs in Capitol Reef Na- tional Park. Rodney L. Player and Philip J. Urness 517 Effects of defoliation on reproduction of a toxic range plant, Zigadentis panicukitus. V. J. Tepedino 524 Distribution and relative abundance of fish in Ruth Reservoir, California, in relation to environmental variables. Steven Vigg and Thomas J. Hassler 529 Temperature and salinity relationships of the Nevadan relict dace. Steven Vigg 541 Observations on woundfin spawning and growth in an outdoor experimental stream. Paul Greger and James E. Deacon 549 Early development of the razorback sucker, Xyrauchen texanus (Abbott). W. L. Minckley and Eric S. Gu.stafson 553 Behavior and habitat preferences of ring-necked pheasants during late winter in cen- tral Utah. Jeffrey G. Skousen and Jack D. Brotherson 562 Rhythm of fecal production and protein content for black-tailed jackrabbits. William D. Steigers, Jr., Jerran T. Flinders, and Susan M. White 567 Prairie dog colony attributes and associated vertebrate species. Tim W. Clark, Thomas M. Campbell III, David G. Socha, and Denise E. Casey 572 Distribution of the moss family Grimmiaceae in Nevada. Matt Lavin 583 Insular biogeography of mammals in the Great Salt Lake. Michael A. Bowers 589 Dorsal hair length and coat color in Abert's squirrel (Sciurus aberti). Denis C. Han- cock, Jr., and Donald J. Nash 597 The raccoon, Procyon lotor, in Wyoming. E. Blake Hart 599 Intercanine crown distances in red foxes and badgers. E. Blake Hart 601 Index 603 ACME BOOKBINDING CO., INC. SEP 9 1983 100 CAMBRIDGE STREET CHARLESTOWN, MASS. 3 2044 072 231 269