HARVARD UNIVERSITY Library of the Museum of Comparative Zoology The Great Basin Naturalist VOLUME 39, 1979 Editor: Stephen L. Wood Published at Brigham Young University, by Brigham Young University TABLE OF CONTENTS Volume 39 Number 1 - March 31, 1979 Phlox longifolia Nutt. (Polemoniaceae) complex of North America. Frederick J. Peab- ody 1 Diatom floristics and succession in a peat bog near Lily Lake, Summit County, Utah. Shobha A. Jatkar, Samuel R. Rushforth, and Jack D. Brotherson 15 Vegetational response to three environmental gradients in the salt playa near Goshen, Utah County, Utah. Michael G. Skougard and Jack D. Brotherson 44 Distribution of sculpins in the Clearwater basin, Idaho. O. Eugene Maughan and Garv E.Saul \. 59 Phytoseiid predators of mite pests in Utah apple orchards. Clive D. Jorgensen and Vich- itra Mongkolprasith 63 Subspecies specificity of gall forms on CJinjsothamniis nauseosus. E. Diirant Mcx\rthur, Charles F. Tiernan, and Bruce L. Welsh 81 Homing by a pygmy rabbit. Jeffrey S. Green and Jerran T. Flinders 88 Beetles from the environs of Lake Powell in southern Utah and northern Arizona. Do- rald M. Allred and Vasco M. Tanner 89 Ants from northern Arizona and southern Utah. Dorald M. Allred and Arthur C. Cole .... 97 Number 2 - June 30, 1979 Review of tularemia in Utah and the Great Basin, Harold E. Stark 103 Chemical composition of some important plants of southeastern Utah summer ranges related to mule deer reproduction. Jordan C. Pederson and K. T. Harper 122 Emergence data and artificial rearing media for an aspen bark beetle, Tn/pophlociis populi (Coleoptera: Scolytidae). David A. Stewart, Gary M. Booth, Jerold L. Petty 129 New synonymy and new species of American bark beetles (Coleoptera: Scolvtidae), Part VIII. Stephen L. Wood '. 133 Annual energy budgets for three common rodent species in the northern Great Basin. R. Kent Sciireiber 143 Preliminary survey of raptor species in the Manti Division, Manti-LaSal National For- est. Stephen G. Jones 155 Vegetation response to a moisture gradient on an ephemeral stream in central Arizona. Deborah Ann Bloss and Jack D. Brotherson 161 Ecological and community relationships of Eriooommi cory))ibosu)n (Polygonaceae) in the Uinta Basin, Utah. Jack D. Brotherson and Karen J. Brotherson 177 Variation in hemoglobin types in the deer mouse {Pcwmyscus mani(ul(itus{ along an al- titudinal gradient. David Wasserman and Donald J. Nash 192 First record of Patapius spiuosiis in Idaho and Nevada (Hemiptera: Leptopodidae). Donald R. Brothers 195 Bee visitation of Phlox hryoides (Polemoniaceae). V. J. Tepedino 197 Ecological distribution oif rodents in Canyonlands National Park, Utah. David M. Arm- strong ' 199 Number 3 - September 30, 1979 igetative and edaphic factors affecting abundance and distribution of small mammals in southeast Oregon. George A. Feldhamer 207 fluence of precipitation and temperature on ring, annual branch increment, and needle growth of White Fir and Douglas-fir in central Utah. John D. Shane and Kimball T. Harper 219 ndies in Nearctic desert sand dune Orthoptera, Part XVI: A new black Stenopelmattis from the Mescalero sands. Ernest R. Tinkham 226 •tential use of Great Salt Lake water for lobster culture. Roger W. Mickelsen, Richard A. Heckmann, and Rex C. Infanger 231 paretic stonefly genera as indicators of ecological parameters (Plecoptera: Insecta). Richard W. Baumann 241 )mpetitive displacement as a factor influencing phytoplankton distribution in Utah Lake, Utah. Lorin E. Squires, Mark C. Whiting, Jack D. Brotherson, and Samuel R. Rushforth 245 -ound nesting and aggressive behavior by the Swainson's Hawk {Btiteo swainsoni). Neil D. Woffinden and James A. Mosher 253 ;sponse of reptile populations to different land management practices on the Idaho National Engineering Laboratory Site. Timothy D. Reynolds 255 screases of juniper woodland in the Utah and Salt Lake valleys since settlement. E. M. Christensen and J. D. Brotherson 263 jmpetition between harvester ants and rodents in the cold desert. Dan S. Landeen, Clive D. Jorgensen, and H. Duane Smith 267 )od habits of burrowing owls in southeastern Idaho. R. L. Gleason and T. H. Craig 274 ora of the Lee Creek valley. Alberta. R. Keith Shaw 277 limates of fescue grasslands of mountains in the western United States. T. Weaver 284 onefly (Plecoptera) records from the basin ranges of Nevada and Utah. Andrew L. Sheldon 289 Number 4 - December 31, 1979 jraginaceae of the southwestern United States. Larry C. Higgins 293 nomiopsyllinae (Siphonaptera: Hystrichopsyllidae), II. The genera Callistopsyllus, Conorhinopsylki, Megarthroglossus, and Stenistomera. Vernon J. Tipton, Harold E. Stark, and John A. Wildie 351 aryotypes of four Artemisia species: A. carruthii, A. filifolia, A. frigida, and A. spin- escens. E. Durant McArthur and C. Lorenzo Pope 419 ariation in leaf anatomy and Cog assimilation in Sitanion hystrix ecotypes. Warren P. Clary 427 idex to Volume 39 433 [HE GREAT BASIN NATURALIS" /olume39 No. 1 March 31, 1979 Brigham Young Universii LIBRARY JUL 23107,3 GREAT BASIN NATURALIST MEMOIRS Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agri- cultural Sciences; Ernest L. Olson, Director, Brigham Young University Press, Univer- sity Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In general, only previously unpublished manuscripts of less than 100 printed pages in length and pertaining to the biological natural history of western North America are ac- cepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natural history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manu- scripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back numbers should be directed to Brigham Young University, Life Science Museum, Pro- vo, Utah 84602. The Great Basin Naturalist Memoirs may be purchased from the same of- fice at the rate indicated on the inside of the back cover of either journal. Scholarly Exchanges. Libraries or other organizations interested in obtaining either journal through a continuing exchange of scholarly publications should contact the Brigham Young University Exchange Librarian, Harold B. Lee Library, Provo, Utah 84602. Manuscripts. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back cover. 11-78 650 34941 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 39 March 31, 1979 No. 1 PHLOX LONGIFOLIA NUTT. (POLEMONIACEAE) COMPLEX OF NORTH AMERICA' Frederick J. Peabody- .\bstract.— Over 1,000 herbarium specimens including 24 type specimens were examined in an attempt to achieve a clearer understanding of the Phlox longifolia complex. Four variables were measured for each specimen and the data were statistically analyzed by discriminant analysis. Using a previously published system of classifica- tion for the complex, approximately 73 percent of the measured variation among the specimens was accounted for by that system. The clustering patterns produced in this first analysis indicated that a more conservative approach would be advisable. The formulation and subsequent application of a modified system produced a grouping of speci- mens that accounted for 95 percent of the measured variation for the four morphological characters considered. From four species and seven subspecies previously recognized, one species with five varieties is proposed. The typi- fication of Phlox longifolia Nutt. and other related taxa is resolved. The classification of plants belonging to the Phlox longifolia Nutt. complex has been fraught with difficulties from the very begin- ning of the nomenclatural history of the group. Infraspecific taxa have been attri- buted to one species and then another, new species have been proposed to deal with the great range of diversity within the group, and confusion has arisen as to the level or rank of recognition best suited for any one entity. The present study has two basic objectives: first, to correlate recent collections and cur- rent descriptions with the type material; and second, to review the classification of the complex in light of a broad sample. The method proposed for the solution of the problem is twofold: A test of the most re- cently proposed classification system (Wherry 1955), and then a similar test of a modified system proposed by me. Statistical analyses were conducted with the use of the IBM 360 computer using the SPSS V602 pro- gram for discriminant analysis from the Sta- tistical Package for the Social Sciences avail- able at Brigham Young University Computer Center. Over 1,000 specimens from nine western United States herbaria were exam- ined and included in the sample. Twenty-four type specimens from an additional 13 her- baria were also examined. Classical methods of taxonomic research were employed in re- viewing type material and adjusting the no- menclature in order to bring the treatment of this complex into agreement with the Inter- national Code of Botanical Nomenclature (Stafleu et al. 1972). The variables tested were the following: vestiture of the inflorescence herbage, corolla tube length, leaf length, and leaf width. Over the past 160 years of taxonomic history these four morphological features have proved to be adequate for separation into infraspecific taxa. They are also easily measured from dried and mounted herbarium specimens. Following the descriptions of each taxon in the taxonomy section is a citation of repre- 'A thesis submitted in partial completion for the degree Master of Science. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602; present address: Department of Botany and PUnt Pathology, Iowa State University, Ames, Iowa 50010. 2 Great Basin Naturalist Vol. 39, No. 1 sentative material examined, followed by a number indicating the total number of speci- mens examined and placed within that taxon. The standard abbreviations of herbaria are those of Holmgren and Keuken (1974). Type specimens were examined and photographed and the photographs deposited in the Herba- rium of Brigham Young University (BRY); they are indicated by an asterisk (°) following the herbarium symbol designation in the list of synonyms. The standard abbreviations of serial publications are those of the Torrey Botanical Club (1969). The standard abbre- viations of separate works are those of Law- rence (1968). Statistical information is avail- able from the author upon request. Acknowledgments The author wishes to acknowledge the many herbaria and their staffs for the loan of material for examination. Appreciation is also extended to Dr. Stanley L. Welsh for valu- able assistance in research and preparation. The constant support of Robyne, my wife, is deeply appreciated. Nomenclatural History Thomas Nuttall (1834) described Phlox longifolia from material gathered by Wyeth in 1833 from the "valleys of the Rocky Mountains generally". Heller (1897) elevated Torrey 's (1859) P. speciosa var. ? stanshuryi to species rank in a cryptic note which af- fords little information as to its salient fea- tures. Subsequent workers influencing this complex include: E. Nelson (1899), Brand (1907), Jones (1895, 1908), A. Nelson (1909, 1912, 1924, 1931), Wherry (1938, 1939, 1940, 1941, 1942, 1943, 1944, 1955, 1956), Peck (1941), Jepson (1943), and Mason (1951). Over the past 163 years, 16 species and 36 in- fraspecific taxa have been named within this complex. Of the 52 names proposed in this group only 5 are recognized as valid in the present study. Taxonomy of Phlox Longifolia Complex Taprooted perennials, 0.3-5 dm tall, from a lignous base; leaves cauline, opposite, sometimes alternate above, 5-70 (100) mm long, 1-7 mm wide, linear to oblanceolate; upper cauline leaves glabrous, pubescent or glandular-pubescent; lower cauline leaves glabrescent, basally connate or distinct; pedi- cels (5) 10-40 (50) mm long, glabrous, pub- escent, or glandular-pubescent; calyx of 5 aristate to cuspidate, basally connate sepals with prominent herbaceous midribs, glabrous except on adaxial surface, pubescent or glandular-pubescent, distinct chartaceous in- tercostal membranes either plicate or flat; corolla tube (8) 10-28 mm long, usually glabrous but sometimes sparsely pilose with- out, the limb 5-15 mm long, incisorate, erose, or entire; stamens imequally inserted on the corolla tube, included or scarsely excerted; style 3-cleft, the ovary 3-loculed, the seeds 1 (2-4) per locule. 2(1). Key to Varieties Inflorescence herbage glabrous except calyx within, or pubescent on upper leaf margins, pedicels and calyx without; leaves linear P. longifolia var. longifolia Inflorescence herbage glandular-pubescent, rarely simply pubescent (var. longipes and var. viridis); leaves linear to oblanceolate 2 Corolla tube 22-28 mm long; southwestern New Mexico westward across Arizona to southern and western Utah, Nevada and eastern California P. longifolia var. stanshuryi Corolla tube 12-19 mm long; variously distributed throughout western North America 3 March 1979 Peabody: North American Phlox 3(2). Leaves (even upper) distinctly oblanceolate, 3-5 mm wide; inflorescence herbage glandular-pubescent; internodes well spaced; stems stout and often trailing; southwestern South Dakota westward to eastern California P. longifolia var. brevifolia Leaves linear to linear-lanceolate, 1-3 mm wide; inflorescence herbage rarely nonglandular-pubescent; internodes well spaced or congested; stems slender and usually ascending; variously distributed throughout western North America 4 4(3). Leaves 45-70 mm long, 2.5-4 mm wide; internodes well spaced; northern New Mexico and Arizona through Nevada, Utah, and western Colorado, narrowly extending into southern Idaho and Oregon P. longifolia var. longipes Leaves 16-35 mm long, 1-2 mm wide; internodes usually congested; Wash- ington, Oregon, Idaho, extending narrowly into extreme northern Utah and Nevada P. longifolia var. viridis Phlox longifolia Nutt. var. longifolia Phlox longifolia Nutt., J. Acad. Nat. Sci. Philadelphia 7: 41. 1834. Holotype: Valleys of the Rocky Mts. generally, 1833, Wyeth s.n. (BM°). Isotype (K"). P. humilis Dougl. ex Hook., Fl. Boreali-Amer. 2: 72. 1838. Holotype: Oregon; barren sandy plains of the Columbia, 1826, Douglas s.n. (K"). P. speciosa Pursh var. B Dougl. ex Hook., Fl. Boreali- Amer. 2: 72. 1838. Holotype: Oregon; on the summit of the Blue Mountains and subalpine range of the Rockies near perpetual snow, no date, Douglas s.n. (K°). P. sabini Hook, pro syn., Fl. Boreali-Amer. 2: 72. 1838. P. speciosa Pursh var. linearifolia Hook., Hooker's J. Bot. Kew Card. Misc. 3: 289. 1851. Holotype: Valley of the Kooskooskie River and adjoining plains, no date, Douglas s.n. (K"). P. linearifolia (Hook.) A. Gray, Proc. Amer. Acad. Arts 8: 255. 1870. P. longifolia Nutt. f. humilis (Dougl. ex Hook.) Voss, Vil- morin Blumengartn 1: 681. 1894. P. longifolia Nutt. ssp. marginata Brand, Das Pflanzenreich 4''": 65. 1907. Holotype: Oregon; steep grassy slopes near Snake River, where it is common, 23 May 1901, Cu.sick 2517, (G°). P. longifolia Nutt. ssp. marginata Brand var. humilis (Dougl. ex Hook.) Brand, Das Pflanzenreich 4^"": 66. 1907. P. longifolia Nutt. ssp. linearifolia (Hook.) Brand, Das Pflanzenreich 4^'": 66. 1907. P. patula A. Nelson, Univ. Wyoming Publ. Sci., Bot. 93: 47. 1924. Holotype: Colorado; Platte Canyon, 19 May 1894, A. Nelson 1589, (RM°). P. marginata (Brand) A. Nelson, Amer. J. Bot. 18: 434. 1931. P. cortezana A. Nelson, Amer. J. Bot. 18: 434. 1931. Holotype: Colorado; Montezuma Co., roadside between Cortez and Mesa Verde National Park, 11 May 1925, A. Nelson 10436 (RW). P. longifolia Nutt. ssp. humilis (Dougl. ex Hook.) Wherry, Proc. Acad. Nat. Sci. Philadelphia 90: 135. 1938. P. longifolia Nutt. ssp. calva Wherry, Proc. Acad. Nat. Sci. Philadelphia 90: 136. 1938. Holotype: Idaho; Butte Co., 13 miles by road southwest of Darling- ton (43°41i/4'; 113°34V2'), 21 June 1931. E. T. Wherry s.n. (PH°). P. longifolia Nutt. ssp. typica Wherry, pro. typ. Notul. Nat. Acad. Nat. Sci. Philadelphia 87: 5. 1941. P. longifolia Nutt. ssp. cortezana (A. Nelson) Wherry, Notul. Nat. Acad. Nat. Sci. Philadelphia 87: 5. 1941. P. grahamii Wherry, Brittonia 5: 60. 1943. Holotype: Utah; Uinta Co., talus slopes west side of Green River, south of mouth of Sand Wash, 4500 ft., elevation, 27 May 1923, Graham 7884 (CM°). P. longifolia Nutt. ssp. a-longifolia Wherry, pro. typ. Morris Arb. Bull. 3: 90. 1956. Plants short to tall, 0.5-5.0 dm; internodes more or less congested or well spaced; leaves opposite, upper leaves glabrous or pubescent, nonglandular, (10) 20-50 (60) mm long, 1.0-2.5 mm wide, linear to linear-lanceolate; calyx glabrous or pubescent with intercostal membranes either plicate or flat; corolla tube 12-16 mm long (Fig. 1). Representative material.— Huntley 889 (WTU); Parker 578 (OSC); Davis 334 (IDS); Vickery 550 (ARIZ); Cronquist 6237 (COLO); Thompson 11316 (MONTU); Brown 3780 (UNM). 645. Distribution.— Central and eastern Wash- ington, eastern and central Oregon, southern Idaho, southwestern Montana, southwestern Wyoming, Nevada, Utah, western Colorado, Great Basin Naturalist Vol. 39, No. 1 Fig. 1. Phlox hngifolia Nuttal var. longifolia. Isotvpe: Valleys of the Rocky Mts. generally. Wyeth s.n., no date (BMr). Measure bar equals 1 cm. March 1979 Peabody: North American Phlox extreme northwestern New Mexico, northern Arizona, and extreme southeastern Cahfor- nia. This nonglandular form is the most abun- dant variety in northern and mesic locahties (Fig. 2). Notes- Nuttall (1834) described Phlox longifoUa based on a specimen with long leaves as compared to the caespitose repre- sentatives of the genus, sic. P. caespitosa, P. hoodii etc. Douglas (1838) described P. hii- milis based on material similar to that of Nut- tail's P. longifoUa, selecting this epithet be- cause the leaves were smaller than those of P. speciosa already described by Pursh (1814) and Lindley (1830). Gray (1870) rectified the situation by placing Douglas's P. humilis in synonymy with Nuttall's P. longifoUa. At the same time Gray proposed a new species which had longer leaves and distinct replica- tion of the intercostal membranes of the ca- lyx, naming this species P. linearifolia (Hook.) A. Gray. With the passage of years since the work of Gray (1870) and the activity of post-Gray- an monographers, the taxonomy of this group became extremely obscured. Nuttall's type was disregarded, Gray's Phlox linearifolia fell out of use and bi- or trinomials were applied to erroneous taxa. The longer-leaved and tall- er P. linearifolia became commonly known as P. longifoUa, leaving the type of P. longifoUa to be erroneously renamed at various in- fraspecific levels. Among these renamings are: P. longifoUa ssp. marginata var. humilis (Dougl. ex Hook.) Brand (1907), P longifoUa ssp. humilis (Dougl. ex. Hook.) Wherry (1938), and P. longifoUa f. humilis (Dougl. ex Hook.) Voss (1894). This unfortunate condi- Fig. 2. Distribution of Phlox longifoUa Nutt. var. longifoUa over western North America. 6 Great Basin Naturalist Vol. 39, No. 1 tion has continued to the present day, caus- ing taxonomic confusion within the complex. Wherry (1943) proposed Phlox grahamii Wherry as an intermediate form between two genera (Phlox and Microsteris). His rea- sons were that the plant in question (Graham 7884) combines morphological characters of each genus. It is perennial and rather tall with well-spaced intemodes and a distinct corolla tube like Phlox. The leaves, however, are relatively small, few, and some are lobed near the base; the corolla tube is strongly flaring at the base and the inflorescence is sparse as in the genus Microsteris. Upon examination of this specimen, which unfortunately is the only specimen of this species ever collected or reported, I have found that it is copiously infested with a rust (Puccinia plumbaria Peck) according to iden- tification made with Arthur (1962). The base of the corolla tube is filled with hyphae, causing it to flare, and one of the few upper leaves bears a number of aecia imbedded in its epidermis that caused it to lobe abnor- mally as it elongated in growth. The general health of the plant was obviously very poor and all of the features noted by Wherry as re- sembling the genus Microsteris are, in my opinion, the result of teratology. Phlox grahamii appears to be a diseased P. long- ifolia var. longifolia and is therefore reduced to synonymy. P. stansburyi (Torr.) Heller ssp. superba (Brand) Wherry, Notul. Nat. Acad. Sci. Philadelphia 113: 4. 1942. Plants moderately tall, 1.5-4.0 dm, inter- nodes well-spaced, 3-6 cm long; leaves oppo- site, upper leaves somewhat glandular-pub- escent, 22-39 mm long, 2.1-3.8 mm wide, linear to linear-lanceolate; calyx glandular- pubescent with the intercostal membranes usually flat; corolla tube 22-28 mm long (Fig. 3). Representative material— Arnoff 37, (WTU), Maguire 25123 (OSC), Ferris 8054 (MONTU), Hershey 2826 (UNM), Wooton s.n. (COLO), Wooton s.n. (ARIZ). 38. Distribution— Western New Mexico, Ari- zona, western Utah, Nevada, and extreme eastern California (Fig. 4). Notes— The taxon stansburyi proves to fit into P. longifolia quite well because it has many characters in common with other vari- eties. The long corolla tube usually cited as the distinguishing feature merely represents a point in a continuum of corolla tube length within this complex. Other features are essen- tially the same as in P. longifolia [sensu lato). As defined here var. stansburyi occurs not only in southwestern New Mexico and adja- cent Arizona (Wherry 1956) but also in northern Arizona, Utah, and Nevada. As it passes northward it appears to assume some of the characteristics of other varieties, most noticeably reduction in leaf size and length of the corolla tube. Phlox longifolia Nutt. var. stansburyi (Torr.) A. Gray Phlox longifolia Nutt. var. stansburyi (Torr.) A. Gray, Proc. Am. Acad. Arts 8:255. 1870. P. speciosa Pursh var. ? stansburyi Torr., Rep. U.S. Mex. Bound. Surv. 2: 145. 18.59. Holotype: New Mexi- co; Dona Ana Co., gravelly hills near the Organ Mountains, .30 April 1852, Bigebw s.n. (NY). P. stansburyi (Torr.) Heller, Bull. Torrey Bot. Club 24: 478. 1897. P. longituba Heller, Muhlenbergia 2: 228. 1906. Isotype (?): California; Inyo Co., Sierra foothills west of Bishop in coarse granite sand, 23 May 1906, Hel- ler 8320 (BW). P. stansburyi (Torr.) Heller ssp. eu-stansburyi Brand, Das Pflanzenreich 4"": 66. 1907. P. superba Brand, Das Pflanzenreich 4"": 67. 1907. Holo- type: Nevada; Nye Co., Tonopah, May 1905, Brown s.n. (UC). P. stansburyi (Torr.) Heller ssp. eu-stansburyi Brand f. longituba (Heller) Wherry, Notul. Nat. Acad. Nat. Sci. Philadelphia 113: 4. 1942. Phlox longifolia Nutt. var. brevifolia (A. Gray) A. Gray Phlox longifolia Nutt. var. brevifolia (A. Grav) A. Gray, Synop. Fl. N. Amer. Vol. 2 Pt. 1: 133. 1878. P. longifolia Nutt. var. stansburyi (Torr.) .\. Gray f. brevifolia A. Gray, Proc. Amer. Acad. Arts 8: 255. 1870. Lectotype: Nevada; Ormsby Co., near Car- son City, 1865. Anderson s.n. (GH"). P. longifolia Nutt. var. stansburyi (Torr.) A. Gray subvar. brevifolia (A. Gray) Watson, C. King, Report of the Geographical Expl. 40th Parallel 5: 261. 1871. P. stansburyi (Torr.) Heller var. brevifolia (A. Gray) E. Nelson, Wyoming Agric. Exp. Sta. Annual Rep. 9: 27. 1899.' P. stansburyi (Torr.) Heller ssp. eu-stansburyi Brand var. brevifolia (A. Gray) Brand, Das Pflanzenreich 4"": 66. 1907. P. grayi Wooton & Standley, pro. syn. Contr. U.S. Nat. Herb. 16: 161. 1913. P. longifolia Nutt. ssp. brevifolia (A. Gray) H. Mason, Abrams 111. Fl. Pac. States 3: 409. 1951. March 1979 Peabody: North American Phlox .0 'Ci, A Fig. 3. Phlox longifolia Nutt. var. stansbtiryi (Torr.) Gray. Topotype: New Mexico, Dona Ana Co., in the Organ Mountains. E. O. Wooton s.n. 28 May 1905 (WTU°). Measure bar equals 1 cm. 8 Great Basin Naturalist Vol. 39, No. 1 Plants shorts, often trailing, 1-3 dm tall, internodes well spaced, 1-4 (5) cm long; leaves opposite, upper leaves glandular-pub- escent, 16-30 mm long, 3-5 (6) mm wide, distinctly oblanceolate; upper sometimes lin- ear-lanceolate; calyx glandular-pubescent, in- tercostal membranes usually flat; corolla tube 13-15 mm long (Fig. 5). Representative material.— Cronquist 8894 (WTU), Lenz 21826 (OSC), Forewood s.n. (K), Welsh 9689 (BRY), Christensen s.n. (BRY), Blauer 7 (BRY), Palmer 308 (GH), Anderson s.n. (GH). 67. Distribution.— Extreme southwest South Dakota, central western Colorado, northern Arizona, Utah, Nevada, and extreme eastern California (Fig. 6). Notes.— Variety brevifolia has been named under innumerable combinations over its tax- onomic history. Gray's (1870, 1878) and Wat- son's (1871) transference of this taxon from one rank to another betrays its complexity. Of all varietal names applied over the past years, Gray's var. brevifolia antedates all and is chosen as the legitimate name. Because Gray indicated no type in the publication of this variety, a lectotype (Wherry 1955) has been designated (the reader is referred to the list of synonyms). It is estimated that the range of this taxon extends further eastward than any other within this complex, crossing the Continental Divide through Wyoming and into the Black Hills of southwestern South Dakota (fide Forewood 1888). Phlox longifolia Nutt. var. longipes (M. E. Jones) M. E. Peck Phlox longifolia Nutt. var. longipes (M. E. Jones) M. E. Peck, Mann. Higher PLS. Oregon 571. 1941. P. linearifolia (Hook.) A. Gray var. longipes M. E. Jones, Contr. W. Bot. 12: 53. 1908. Holotype: Idaho; Fig. 4. Distribution of Phlox longifolia var. stanshuryi (Torr.) Gray over western North America. March 1979 Peabody: North American Phlox Washington Co., Weiser, 28 April 1900, Jones s.n. (POM°). P. longifolia Nutt. var. puberula E. Nelson, Wyoming Agric. Exp. Sta. Annual Rep. 9: 26. 1899. Holo- type: Wyoming; Uinta Co., Evanston, 5 June 1898, A. Nelson 4544 (RM°). P. viscida E. Nelson, Wyoming Agric. Exp. Sta. Annual Rep. 9: 25. 1899. Holotype: Oregon; Columbia Co., Blue Mountains, 15 July 1896, Piper 2397 (US'). P. stansburyi (Torr.) Heller ssp. compacta Brand var. vis- cida (E. Nelson) Brand, Das Pflanzenreich 4"°: 67. 1907. P. stansburyi (Torr.) Heller ssp. compacta Brand var. puberula (E. Nelson) Brand, Das Pflanzenreich 4^"^": 67. 1907. P. stansburyi (Torr.) Heller ssp. eu-stansburyi Brand var. brevifolia (A. Gray) Brand subvar. microcalyx Brand, Das Pflanzenreich 4"": 67. 1907. Holo- type: Arizona; Yavapai Co., Prescott Mountain District, 1876, Farmer 397 (G*). P. puberula (E. Nelson) A. Nelson, Manual Bot. Rocky Mts.: 397. 1909. P. longifolia Nutt. var. filifolia A. Nelson. Bot. Baz. (Crawfordsville) 54: 143. 1912. Holotype: Idaho; Blaine Co., Ketchum midst sagebrush, stream bottoms, alt. 5887, 19 July 1911, A. Nelson 1192 (KM"). P. longifolia Nutt. ssp. compacta (Brand) Wherry, Proc. Acad. Nat. Sci. Philadelphia 93: 135. 1938. P. longifolia Nutt. ssp. longipes (M. E. Jones) Wherry, Proc. Acad. Nat. Sci. Philadelphia 93: 135. 1938. P. viridis E. Nelson ssp. longipes (M. E. Jones) Wherry, Morris Arb. Bull. 3: 88. 1955. P. viridis E. Nelson ssp. compacta (Brand) Wherry, Bail- eya 4: 98. 1956. Plants moderately tall 1.5-4.0 dm, inter- nodes somewhat congested or well-spaced 1-5 cm long; leaves opposite, upper leaves glandular-pubescent, 3-10 (15) mm long, 1.7-3.6 mm wide, linear to linear-lanceolate; calyx glandular-pubescent with the inter- costal membranes usually flat; corolla tube 14-18 mm long (Fig. 7). Representative material.— Whites 1037 (WTU), Peck 7790 (OSC), Davis 3082 (IDS), Cottam 5062 (ARIZ), Clokey 7630 (MONTU), Clark s.n. (UNM), Hitchcock 20432 (COLO). 122. Distribution — Eastern Oregon, southern Idaho, Utah, western and southern Colorado, Nevada, northern Arizona, northwestern Fig. 5. Phlox longifolia Nutt. var. brevifolia Gray. Collection: South Dakota, Black Hills. W. H. Forwood 1887 (K°). Measure bar equals 1 cm. 10 Great Basin Naturalist Vol. 39, No. 1 New Mexico, and extreme eastern California (Fig. 8). Notes,— Even though this taxon is found in northern as well as southern areas it appears to be more abundant in southern Utah, Ari- zona, New Mexico, and southwestern Colo- rado. The long, narrow leaves and glandular pubescence are distinctive. It appears that var. longipes may be a transitional form be- tween the pubescent var. longifolia and the glandular-pubescent var. stansburyi. The overlapping pattern in distribution and mor- phology would point to this possibility. Phlox longifolia Nutt. var. viridis (E. Nelson) Peabody stat. nov. Phlox viridis E. Nelson, Wyoming Agric. Exp. Sta. An- nual Rep. 9: 25. 1899. Holotype: Washington; Kittitas Co., Ellensburg, 20 May 1897, Piper 2689 (WS-). P. stansburyi (Torr.) Heller ssp. compacta Brand var. puberula (E. Nelson) Brand subvar. viridis (E. Nelson) Brand, Das Pflanzenreich 4"°: 67. 1907. P. longifolia Nutt. ssp. viridis (E. Nelson) Wherry, Notul. Nat. Acad. Nat. Sci. Philadelphia 87: 5. 1941. P. viridis E. Nelson ssp. a-viridis Wherry, pro. typ. Mor- ris Arb. Bull. 3: 88. 1955. Plants short 1-3 dm, stem much branched from the base, internodes 1-2 cm long; leaves opposite, upper leaves glandular-pubescent, 16-30 mm long, 1.0-1.5 mm wide, linear; ca- lyx glandular-pubescent, intercostal mem- branes usually flat; corolla tube 13-15 mm long (Fig. 9). Representative material.— Hitchcock 17409 (WTU), Peck 25922 (OSC), Davis 99- 36 (IDS), Stevens 150 (BRY), Stevens 161 (BRY), Piatt 157 (BRY). 142. Distribution.— Central Washington, cen- tral and southeastern Oregon, southern Fig. 6. Distribution of Phlox longifolia Nutt. var. brevifolia Gray over western North America. March 1979 Peabody: North American Phlox 11 Idaho, extreme northern Utah and Nevada (Fig. 10). Notes.— Easily distinguished by its fine, narrow leaves, multiple branches, and usually congested intemodes, var. viridis is primarily of northern distribution, extending southward along the western slope of the Rocky Moun- tains into northern Utah and Nevada. Literature Cited Abrams, L. 1954. Illustrated flora of the Pacific States. Vol. 4, Stanford University Press, Stanford, Cali- fornia. Arthur, J. D. 1962. Manual of the rusts of the United States and Canada. Hafner Publishing Co., New York. 4,38 pp. Bentham, G. 1845. Polemoniaceae. In: A. de Candolle, Prodromus 9: 302-322. Brand, A. 1907. Polemoniaceae. In: Das Pflanzenreich 4^^": 1-203. Fig. 7. Phlox hngifolia Nutt. var. hngipes (M. E. Jones) Peck. Holotype: Idaho, Washington Co., Weiser. M. Jones s.n. 28 April 1900 (POM*). Measure bar equals 1 cm. 12 Great Basin Naturalist Vol. 39, No. 1 Chaudhri, M. N., I. H. Vegteb, and C. M. DeWal. 1972. Index herbariorum, Part Il-Collectors (E-H). Regnum Vegetable, Vol. 9. Kemink en Zoon N. V. Utrecht, Netherlands. 296-473. Cronquist, a. 1964. In: C. L. Hitchcock, et al. Vascular plants of the Pacific Northwest. Univ. Wash. Publ. Biol. 17(4): 1-510. Fisher, R. A. 1936. The use of multiple measurements in taxonomic problems. Ann. Human Genetics 7: 179-188. Gray, A. 1870. Revision of the North American Polemo- niaceae. Proc. Amer. Acad. Art. 8: 247-282. 1878. Polemoniaceae. Synoptical flora of North America, Vol. II, Pt. 1. John Wilson and Sons, Cambridge, Massachusetts. Heller, A. A. 1897. Notes on plants of New Mexico. Bull. Torrey Bot. Club 24: 477-480. 1906. Polemoniaceae. Muhlenbergia 2: 228-235. Holmgren, P. K., and W. Keuken. 1974. Index her- bariorum, Pt. I, The herbaria of the world. Reg- num vegatabile. Vol. 92. Dosthoek, Scheltema and Holkema, Utrecht, Netherlands. 397 pp. Hooker, W. J. 1829-1834. Flora Boreali-Americana. Vol. I. G. Bohn, London. 298 pp. 1851. Catalogue of Mr. Geyer's plants collected in the upper Missouri. Hooker's J. Bot. Kew Card. Misc. 3: 273-305. Jepson, W. L. 1925. Manual of the flowering plants of California. .Associated Student Stores, Berkeley, California. 1238 pp. 1943. A flora of California, Vol. 3. University of California Press, Berkeley, California. 284 pp. Jones, M. E. 1895. Contributions to western botany. Contr. W. Bot. 8: 1-43. 1908. Contributions to western botany, No. 12. Contr. W. Bot. 12: 1-100. Kuntze, O. 1891. Rev. Gen. Bot. Pars. 2. H. Sturtz, Wursburg. Lanjouw, J., and F. a. Stafleu. 1954. Index herbario- rum. Part II, Collectors (A-D). Regnum Vegeta- bile, Vol. 2. Kemink en Zoon N. V., Utrecht, Netherlands. 1-174. 1957. Index herbariorum. Part II, Collectors (E-H). Regnum vegatabile, Vol. 9. Kemink en Zoon N.V., Utrecht, Netherlands. 175-295. Lawrence, G. M. H., ed. 1968. Botanico-Periodicum- Huntianum. S-H Service Agency, Inc., New York, 1063 pp. Fig. 8. Distribution oi Phlox hngifolia Nutt. var. longipes (M. E. Jones) Peck over western North America. March 1979 Peabody: North American Phlox 13 LiNDLEY, J. 1830. "Phlox speciosa." In: Edward's Bot. Reg. 16: pi. 1.351. Mason, H. L. 1925. Abrams illustrated flora of the Pacif- ic States. Stanford University Press, Stanford, California. Nelson, A. 1909. "Phlox puberula." In' Nelson, A. and J. M. Coulter, New manual of botany of the central Rocky Mountains (vascular plants). American Book Co., New York. 646 pp. 1912. "Phlox longifolia var. filifolia." In: Bot. Gaz. (Crawfordsville) 54: 143. 1924. Taxonomic studies by Aven Nelson I. Phloxes, new and old. Univ. Wyoming Publ. Sci. Bot. 1(93): 47-68. 1931. New species from mountains and deserts. Amer. J. Bot. 18: 431-441. 1931. Phlox longifolia and Phlox stansburyi and their immediate relatives. Amer. J. Bot. 18: 441-442. Nelson, E. 1899. Revision of the western North Ameri- can Phloxes. Wyoming Agric. exp. Sta. Annual Rep., No. 9: 1-36. NiE, N. E., ed. 1975. Statistical package for the social sciences. 2d ed. McGraw-Hill Book Co., New York. NuTTALL, T. 1834. Plants of the Rocky Mountains. J. Acad. Nat. Sci. Philadelphia 7: 5-60. Peck, M. E. 1941. A manual of the higher plants of Ore- gon. Binfords and Mort, Portland, Oregon. 866 pp. 1941. Validation of new combinations. Madrono 6: 135-1.36. Fig. 9. Phlox longifolia Nutt. var. viridis (E. Nelson) Peabody. Holotype: Washington, Kittitas Co., Ellensburg. C. V. Piper 2689 20 May 1897 (WS°). Measure bar equals 1 cm. 14 Great Basin Naturalist Vol. 39, No. 1 PuRSH, F. 1814. Flora Americae-Septentionalis. In two volumes. White, Cochrane and Co., London. Stafleu, F. a. 1967. Taxonomic literature. Regnum veg- etabile. Vol. 52. Inter-documentation Co. AG, Utrecht, Netherlands, 566 pp. 1972. International code of botanical nomencla- ture. Regnum vegetabile. Vol. 82. A. Oosthoek's Uitgeversmaatschappij N. V., Utrecht, Nether- lands. 426 pp. ToRREY Botanical Club. 1969. Index to American bot- anical literature. In four volumes. G. K. Hall and Co., Boston, Massachusetts. ToRREY, J. 1859. Botany of the boundary. In. Emory, Re- port on the United States and Mexican boundary survey. Vol. 2. C. Wendell, Printers, Washington, D.C. Voss, J. 1894. Phlox longifolia f. humilis. In: Vilmorin Blumengarten 1: 681. Watson, S. 1871. The Botany. In: C. King, Report of the geographical exploration of the fortieth paral- lel. Vol. 5. U.S. Government Printing Office, Washington, D.C. Wherry, E. T. 1938. Phloxes of Oregon. Proc. Acad. Nat. Sci. Philadelphia 90: 133-140. 1939. Four southwestern subspecies of Phlox. J. Wash. Acad. Sci. 29: 517-519. 1940. Geographic relations in the genus Phlox. Bartonia 20: 12-14. 1941. Phloxes of Idaho. Notul. Nat. Acad. Nat. Sci. Philadelphia 87: 5-13. 1940. A provisional key to the Polemoniaceae. Bartonia 20: 14-17. 1942. Phloxes of Nevada. Notul. Nat. Acad. Nat. Sci. Philadelphia 113:4-11. 1943. Microsteris, Phlox, and an intermediate. Brittonia 5: 60-63. 1944. New Phloxes from the Rocky Mountains and neighboring regions. Notul. Nat. Acad. Nat. Sci. Philadelphia 146: 1-11. 1955. The genus Phlox. Morris Arb. Bull. 3: 1-174. 1956. Validation of new combinations in Phlox. Baileya 4: 97-98. WooTON, E. O., AND P. C. Standley. 1913. New Plants from New Mexico— Polemoniaceae. Contr. U.S. Nat. Herb. 16: 160-162. Fig. 10. Distribution of Phhx longifolia Nutt. var. viridis (E. Nelson) Peabody over western North America. DIATOM FLORISTICS AND SUCCESSION IN A PEAT BOG NEAR LILY LAKE, SUMMIT COUNTY, UTAH Shobha A. Jatkar,' Samuel R. Rushforth,' and Jack D. Brotherson' Abstract.— Diatoms from core samples obtained from the edge of Lily Lake, Summit County, Utah, were studied. Populations from each 15 cm of the core and niche metrics of the important species were analyzed. This study dem- onstrates four periods of history in the lake as mirrored by the diatom populations. The first period was mesotrophic and alkaline. This graded into an acidic, dystrophic period which in turn yielded again to a mesotrophic alkaline period. The most recent period can be characterized by a return to dystrophy. Paleodiatomological investigations report- ed to date are generally based on core sam- ples either from marine or lake sediments, and a relatively large literature is extant on this subject. However, fossil diatom studies of bogs are relatively rare. We believe the pres- ent study represents the first detailed account of fossil and living diatoms from bog samples within the Great Basin of western North America. Most bogs in high mountain regions are formed by hydrarch succession following gla- ciation. Such bogs are generally acidic, high- ly humic, and poor in nutrients. Because of these specific features, bog floras are dis- tinctive in biotic composition. In addition, due to high acidity and other factors, bog de- posits often contain fossil or subfossil diatoms in an excellent state of preservation. The first bog study in America was that of Lewis (1863) based on a marshy area around a shallow pond in Notch Valley in the White Mountains of New England. He observed many diatoms and desmids in his samples and drew conclusions on the relationship between the flora and geology of the area. Patrick (1943) studied Linsely Pond, Con- necticut, and observed significant changes in the diatom flora at different depths. The maximum number of diatoms, but with a low species diversity, occurred at a depth of 12 m. Species diversity increased between 12 and 9.6 m. Shallow water or littoral forms oc- curred in sediments from 8.4 to 6 m, but the best developed planktonic flora was observed at 5.1 m. Based on these and other observa- tions she concluded that recent changes in the diatom flora were more or less due to the use of part of the drainage basin for agricul- tural purposes. Weaver (1948) studied the diatom flora from acidic Lakeville Bog, Indiana, where he observed freshwater sponge spicules and no diatoms in sediment layers below 6.9 meters from the surface. He found maximum diatom populations in layers between 4.5 and 3.3 m and very few diatoms between 3.3 m and the surface. He found species of Pinnularia, Gomphonema, and Eunotia indicative of dys- trophy between 4.5 and 3.3 m below the sediment surface. From these observations he inferred that during its early history, the lake was deep, clear, and poor in nutrient content. Certain sponges tolerant of oligotrophic con- ditions grew well in the lake during this time. Later the lake became richer in nutrients and diatoms became more prevalent. It was con- cluded that this lake changed from oligotro- phic to dystrophic during its history. Reimer (1961) studied the diatom flora of alkaline Cabin Creek raised bog, Indiana. He found that centric diatoms were absent and only active motile diatoms were present. Ac- cording to Reimer this was possibly due to the greater adaptability of motile as opposed to nonmotile forms. He found very few Nitz- schia species and an abundance of Cymbella leptoceros. From these and other observations he inferred that Cabin Creek raised bog ex- hibited oligotrophic conditions. Collingsworth, Matthew, and Collins (1967) observed at Vestaburg Bog, Montcalm 'Department of Botany and Range Science, Brigham Young University, Provo, Utah. 15 16 Great Basin Naturalist Vol. 39, No. 1 County, Michigan, a good preservation of di- atoms to 14 m below the bog surface. They found a gradual increase in the diatom popu- lation from a depth of 7 m to the surface and speculated that this lake must have been oli- gotrophic when it originated by glaciation, later becoming eutrophic, and finally dystro- phic. Very few paleodiatomological studies have been made in Utah. Patrick (1936) collected sediment samples from various regions of the Great Salt Lake using a Musselman Peat Sampler. She identified and prepared a list of diatoms collected from each section. Based upon species present, she described the habi- tat of deposition for each core (especially the salinity) and inferred past ecological condi- tions. Patrick inferred that a freshwater lake formerly occupied the region where the Great Salt Lake occurs now. The findings of fresh and brackish water diatoms supported her conclusion that swamps existed here in Pleistocene or early Post-Pleistocene times. Lake Bonneville (the fresh water lake for- merly occupying this region) gradually be- came more and more saline due to evapo- ration and lack of drainage. A marine diatom flora was never established in the lake, even though the lake went through a stage when the salinity was the same as normal marine waters. Patrick found no living diatoms in the most saline parts of the lake. However, she did obtain a few living diatoms from river deltas and areas surrovmding brackish mar- shes where the salt concentration was not as high as in the lake proper. Hasler and Crawford (1938) studied fossil diatoms in samples from Lake Bonneville marl sediments and foimd 16 genera and 27 species of diatoms. These authors drew few ecological conclusions from their data but in- ferred that Lake Bonneville was saline during some portion of the deposition period. Setty (1963) studied fossil diatoms of the Pleistocene lacustrine sediments of Bonne- ville Basin. He found 126 taxa and presented a systematic and paleoecological discussion of these diatoms. Bolland (1974) studied diatom deposits in a single 5 m core from Utah Lake, Utah. He observed 155 diatom taxa and studied succes- sion through the core. By analyzing chemical and successional data, Bolland concluded that Geneva Works of United States Steel, which is located on the east shore of the lake, has had a major effect in bringing about changes in water chemistry of the lake. From these and other observations he inferred that Utah Lake is progressing from a deep water me- sotrophic system to shallow water eutrophic. According to Bolland, this is mainly due to industrial development in the surrounding area and the inflow of large amounts of sew- age into the lake. Several reports have been published on geological, ecological, and floristic surveys of the Uinta Moimtain region of Utah. Atwood (1909) has given an account of glaciation of the Uintas and the formation of many pot- hole lakes in this region. Norrington (1925) surveyed the algal flora of several mountain lakes and streams of the Wasatch and Uinta Ranges and discussed some ecological aspects of this flora. Cottam (1930) surveyed the flora of the Uinta Mountains and discussed some unusual features of this area. The algal flora of Mirror Lake was surveyed by Snow and Stewart (1939). Stutz (1951) studied some Uinta Mountain lakes from a floristic point of view. He performed analyses of pH, moisture content, and organic matter content from several lake sediments. Hay ward (1952) con- ducted ecological studies in the Uinta Moun- tains and listed the characteristic plants of the ponds and surrounding zones. Christensen and Harrison (1961) studied the ecology of flowering plants and gymnosperms at Lily Lake. Vincent (1963-1964) provided infor- mation on the location, area, and depths of 100 Uinta Mountain lakes and included infor- mation on fish productivity of these lakes. Coombs (1964) made a floristic and ecologi- cal survey of the algae of the western Uinta Mountains and adjacent areas. He also gave a short geological account of the area. Palmer (1968) provided a floristic and ecological sur- vey of the algal flora of Lily Lake. Firmage (1969) studied the flora and conifer succes- sion of a few lakes and ponds in the Trial Lake region. Hansen (1971) published a re- view article on the developmental history of the Uinta Moimtains. Lawson and Rushforth (1975) studied the diatom flora of the Provo River, Utah. They found 225 taxa, many of which were observed in the Uinta Mountain region. March 1979 Jatkar et al.: Lily Lake Diatoms 17 The objectives of the present study were twofold: (1) to identify and characterize all fossil and living diatoms from Lily Lake; and (2) to determine if the pattern of diatom suc- cession would illuminate the developmental history of this and similar Uinta Mountain bog lakes. Study Area Our studies were performed on samples collected from an acid Sphagnum bog on the edge of Lily Lake in the Uinta Mountains, Summit County, Utah. Lily Lake is represen- tative of the many Sphagnum moorland lakes in the subalpine zone of these mountains. Moorland lakes, according to Tanner's (1931) system of classification, are characterized by Sphagnum banks, mucky bottom, and a con- stant surface level throughout the year. Lily Lake demonstrates all of these character- istics. Lily Lake is about 34 km northeast of Kamas, Summit County, Utah. It is about 1 km west of the larger Trial Lake at an eleva- tion of 3,280 m. It has a surface area of 1.2 ha and a maximum depth of 5 m (Vincent 1963-1964). Access to this lake is by foot trail from Trial Lake. The annual snow fall in this area is around 189 cm (Whaley and Jones 1977). Snow usu- ally persists until July and therefore the area is inaccessible for up to eight months a year. Lily Lake is surrounded by a sedge meadow which occurs in the peat soil. Spaghnum moss is the most important component of this meadow. Near the shore of the lake, pure stands of Sphagnum are common. A dense growth of the water lily Nuphar polysepahim (Engelm.) Green covers much of the lake sur- face. Lily Lake was selected for the present in- vestigation for several reasons. First, it is in- accessible for much of the year and is away from a main road. In addition, the area has been set aside as a botanical station for Brig- ham Young University. Thus, the area re- mains largely imdisturbed by human activi- ties, thereby avoiding soil erosion and intermixing of the fossil flora. Second, this lake represents a typical acid bog type lake (pH = 4.5 to 5.5) in the Uinta Mountains. It has acidic, organic sediments with low miner- al content, and its water contains low levels of oxygen and nutrients. An excellent preser- vation of diatoms was noted for specimens collected down to 4 m below the bog surface. Third, due to its unique geological and geo- graphical setting the area has been in- fluenced by several factors, including glacia- tion to form lake basins, hydrarch succession to fill such basins, the admixture of southern and northern vascular floristic elements (Cot- tam 1930), etc. The present study was under- taken to shed light on these factors as well as to serve as a taxonomic and ecological dia- tom survey. Geology The Uinta Mountains are in northeastern Utah (Fig. 1). They consist of a single range of peaks, extending west to east from Kamas Prairie, Utah, to the Little Snake River Val- ley in northwestern Colorado. This mountain range is about 240 km long and about 56 km wide. If the crest line of the range continued westward, it would cross the Wasatch range of Utah at nearly a right angle and reach the Great Bonneville Basin a few kilometers south of Salt Lake City. These two ranges are (->'■;* Uinta Mts. ^ U Provo i£ UTAH Fig. 1. Map of Utah showing the location of the Wasatch and Uinta Mountains. 18 Great Basin Naturalist Vol. 39, No. 1 separated by approximately 16 km. The strata of the Uinta Mountains are of about the same composition as the rest of the Cor- dillerean system to which they belong (For- rester 1937). In contrast to the remainder of the Cordillerean system (including the Rock- ies on the east and Pacific ranges on the west) which lie north-south, the Uinta Moun- tain range is oriented east-west. These moun- tains represent the only major east-west mountain chain in North America. The Uinta Mountains are also unique since they show an absence of igneous activity (Emmons 1907). The lowest point of the Uinta Range is 1,816 m above sea level at Hailstone Junc- tion. The highest point is 4,114 m at King's Peak. The range descends very abmptly at its west end and rather gradually on the east. As noted by Cottam (1930) the range has a large radiating surface which forms a highly dis- sected dome of great surface area. The area receives a large amount of precipitation and has relatively high temperature conditions which allow for the growth of good quality timber. Many geologic features are evident in the Uinta Mountains due to active glaciation at different times in the history of the range. The oldest formation exposed is a Pre- Cambrian quartzite which forms the major constituent of the range. This quartzite is represented chiefly by dark red sandstones which become more and more compact to- ward the west. In the western half of the range, dark purplish red quartzite pre- dominates. Interstratified layers of argil- laceous shales and coarse grits are common. In the coarser beds there is considerable ad- mixture of broken feldspar crystals. The ar- gillaceous beds are of greenish or brownish color and are up to 30 m thick in many places. Cambrian rocks (the Tintic Quartzite) lie unconformably over the Pre-Cambrian. Comformably overlying the Tintic Quartzite is the Ophir Shale. A well-defined angular unconformity representing an extensive ero- sional interval occurs between Cambrian rock and the overlying Mississippian lime- stones. The sediments overlying the Mis- sissippian limestones are Pennsylvanian quar- tzites, which in turn are overlain conformably by Park City (Permian) rocks. Triassic rocks in the Uinta sequence are rep- resented by Woodside Shale, the Thaynes Formation, and Ankareh Shale. At the top of the Triassic sequence the Triassic-Jurassic Nugget Sandstone is present. Wherever it is exposed, the cross-bedded, wind-laid Nugget is an excellent horizon marker throughout the Uinta Range. Conformably overlying the Nugget Sandstone is the Twin Creek Forma- tion (Jurassic). The Morrison Formation, ten- tatively placed in the upper Jurassic, is well exposed along the flanks of the range. Cre- taceous Mesaverde, Frontier, and Mancos formations overlie the Jurassic. Wasatch (Eocene) sediments, where exposed, lie un- conformably on the Mesaverde and older for- mations. Along much of the north flank of the range, the Green River and Bridger for- mations (Eocene) are the lowest Tertiary sediments exposed. Duchesne River (Oligo- cene) sediments are not found on the north flank of the range but are prominent along much of the south slope. Pleistocene glacial drift and debris cover much of the underlying structure in the Uinta Range, and it is often difficult to separate these late deposits from older Tertiary sediments. As reported by Atwood (1909) all the larger canyons have a characteristic U-shape due to glaciation. Their upper portions have been well cleaned by the ice, but the middle and lower portions often contain heavy mo- rainic deposits. The floors of the basin in which the ice formed are all above 2,743 m but there is no sign of ice on the flanks of the range even at the 3,048 m level. Atwood (1909) also presented evidence of at least two separate epochs of glaciation. The earlier epoch was presumably the longer because the ice of that epoch was thicker and extended farther down the canyons. About 30 major glaciers occurred in this earlier epoch and about 39 in the later epoch. Glaciation often has a great influence on drainage systems. The hundreds of lakes and marshes in the Uinta Moimtains indicate that the drainage has been greatly modified by the ice. The area is drained by many stream- lets which feed into the Bear River draining northward, the Weber River westward, the Provo River southwest, and the Duchesne River and Rock Creek southeast. Among these mountains, there are now more than 550 glacial lakes (Atwood 1909). Lily Lake is March 1979 Jatkar et al.: Lily Lake Diatoms 19 one of these lakes. There were many more lakes at the close of glaciation, but since then succession has progressed to completion in many to create meadows. Methods Samples were collected in October 1975 from the northeast side of Lily Lake using a peat borer. A 3.76 m core was taken and sub- sampled every 15 cm. Two samples of the living flora were also collected, including one from the water of the lake and the other from the mucky bottom and moss at the edge of the lake. All samples were examined in the field and their color and organic content re- corded. The samples were transferred to clean glass bottles, labeled, and sealed. Samples were brought to the laboratory and refrig- erated until further processing. A small quan- tity of material from each sample was boiled in concentrated nitric acid, and permanent diatom slides were prepared by standard methods (St. Clair and Rushforth 1977). Dia- toms were mounted in Hyrax mounting me- dium. Diatoms were identified under lOOOX oil immersion using a Zeiss RA microscope. Counts of at least 400 diatom frustules were made for all sample depths except for sample Number 1 from the bottom of the core, which contained few diatoms. The absolute and relative density for each species at each sample depth was computed. Species diversity values were also calculated for each sample using the Shannon-Weaver (Shannon and Weaver, 196.3) formula. Re- gression analyses of floral diversity versus sample depth were conducted using standard statistical methods. Similarity indices between all samples in the study were calculated (Ruzicka 1958), and cluster analysis (Sneath and Sokal 1963) based upon similarity indices was performed. The most important species in each of the groups delineated by cluster analysis were - determined using PxF indices (percent fre- quency times average percent density; Warn- er and Harper 1972). A list of prevalent species was prepared by selecting species present in 30 percent or more of the samples. Niche breadth and niche overlap values were obtained (Colwell and Futuyma 1971) for the prevalent species. A cluster analysis based on the niche overlap of the.se species was then performed (Cody 1974). Where possible, the pH preference and saprobien spectrum for the diatoms encoun- tered in this study were determined. This in- formation was tabulated and plotted. Results Discussion Taxonomy The 157 diatom taxa encountered in this study are arranged systematically in the fol- lowing section. Measurements of length and width (or diameter), number of striae, and number of costae are given for each taxon when applicable. A brief discussion of the distribution and maximum relative frequency of each taxon is given. References to detailed descriptions for each taxon are provided. The slides upon which these determina- tions were made have been deposited in the Brigham Young University collections. Coscinodiscaceae Melosira dickiei (Thwait.) Kutzing: Valve diameter 6-15 ; punctae scattered in central region. (Hustedt 1930:86). Frustules were present throughout the core though their percent frequency was low. It constituted 2 percent of the total diatom population from 150 to 135 cm. Melosira distans (Ehr.) Kutzing: Valve di- ameter 12-17 jLim; striae punctate, 10-15 in 10 jum (Hustedt 1930:93). It was scattered in core samples only from 360 to 225 cm. The maximum number of frustules was observed in samples from 360 to 330 cm. Melosira distans var. pfaffiana (Reinsch) Grunow: Valve in girdle view 7 jum long, 6 jum wide, striae 15 in 10 jum. (Hustedt 1930:93). Frustules were very rare and une- venly distributed in the core. Melosira granulata (Ehr.) Ralfs: Valve in girdle view 7-17 jum long, 5-16 jum wide; striae 10-15 in 10 /xm; spines 10 in 10 jtim (Hustedt 1930:87). Though frustules were scattered throughout the core, they were most common from 180 to 75 cm. They ex- 20 Great Basin Naturalist Vol. 39, No. 1 hibited great variation in size and especially in size of punctae. Melosira italica (Ehr.) Kiitzing: Valve in girdle view 10-14 jum long, 10-15 jum wide; striae 18-20 in 10 jum (Hustedt 1930:91). Frustules were present from 360 to 345 cm but only very rarely observed in samples from other depths. It constituted 4.2 percent of the diatom population at 345 cm. Cyclotella antiqua W. Smith: Valve diam- eter 22 jum; striae 14-20 in 10 fxm (Hustedt 1930:102). Frustules were only rarely ob- served and usually broken. A single entire specimen was observed in a sample from 270 cm. Cyclotella catenata Brun: Valve diameter 17 jum; striae about 10 in 10 jum (Hustedt 1930:108). A single frustule was observed at 120 cm. Cyclotella comensis Grunow: Valve diam- eter 8 ]u,m; striae 20 in 10 pun (Hustedt 1930:102). Only one unbroken specimen was observed at 105 cm. Cyclotella kiitzingiana Thwaites: Valve di- ameter 14-15 jum; striae 9-10 in 10 jum (Hus- tedt 1930:98). Frustules were present from 210 to 45 cm. The maximum number was ob- served in samples from 180 cm and 105 cm where they constituted 2.6 percent of the di- atom population. Cyclotella meneghiniana Kiitzing: Valve diameter 8-8.5 /xm; striae 14-20 in 10 jum (Hustedt 1930:100). Frustules were very rare and were observed only in samples from 280 to 255 cm. Their maximum percent frequen- cy was 0.4 percent at 255 cm. Cyclotella striata (Kiitzing) Grunow: Valve diameter 25 jum; striae 8-10 in 10 [xm (Hustedt 1930:101). A single specimen was observed at 150 cm. Stephanodiscus astrea var. minutula (Kiitz.) Grunow: Valve diameter 12-18 jum; striae up to 15 in 10 jum (Hustedt 1930:110). It was observed in samples from 360 to 135 cm. The maximum number of frustules (1.2 percent) was observed at 180 cm. Stephanodiscus dubius (Fricke) Hustedt: Valve diameter 28-35 jum; striae 8-12 in 10 urn (Hustedt 1930:109). Frustules were pres- ent from 180 to 90 cm. The maximum num- ber was observed in samples from 135 cm, where they constituted 2 percent of the dia- tom population. Stephanodiscus niagarae (Ehr,) Grunow: Valve diameter 28-38 jum; striae 10-15 in 10 jLtm (Cleve-Euler 1951:53). Frustules were very rarely observed. The maximum number was found in samples from 180 cm, where they constituted 0.6 percent of the diatom population. Coscinodiscus odontodiscus Grunow: Valve diameter 26-27 jum; striae 16-17 in 10 jum (Cleve-Euler 1951:59). Frustules were ob- served from 210 to 105 cm. The maximum relative density was 0.6 percent in samples from 120 cm. Coscinodiscus rothii (Ehr.) Grunow: Valve diameter 27 ]u.m; aerolae about 16 in 10 jum (Hustedt 1930:112). A single specimen was observed at 120 cm. Fragilariaceae Tabellaria fenestrata (Lyngb.) Kiitzing: Valve 25-51 jum long; 5-7 /xm wide; striae 20-22 in 10 jum (Hustedt 1930:122). It was observed very frequently in samples from throughout the core. The maximum relative density was 25.7 percent at 225 cm and the lowest was 0.2 percent at 105 cm. Tabellaria flocculosa (Roth) Kiitzing: Valve 13-17 [xm long; 6-8 jum wide; striae about 18 in 10 jum (Hustedt 1930:123). Frus- tules were observed from 210 to 30 cm and also in surface samples. The maximum rela- tive density was 5.6 percent at 30 cm. Diatoma anceps (Ehr.) Kirchner: Valve 24 jLim long by 4-5.5 jum wide; costae 5 in 10 jum; striae 15-20 in 10 jum (Patrick and Rei- mer 1966:106). It was observed as an entire specimen only once at 270 cm. Diatoma hiemale var. mesodon (Ehr.) Grunow: Valve 14-15 jum long by 7-8 jum wide; costae 3-4 in 10 jum; striae about 18 in 10 jum (Patrick and Reimer 1966:108). Frus- tules were very rarely observed in scattered samples in the core. Diatom vulgare Bory: Valve 43-48 jum long by 12-14 /xm wide; costae 6-8 in 10 jtxm; striae about 14 in 10 jtxm (Patrick and Reimer 1966:109). It was observed in samples from 180 to 120 cm. The maximum occurrence was from 180 cm to 150 cm, where it con- stituted 3.6 percent and 5.2 percent, respec- tively, of the total diatom population. March 1979 Jatkar et al.: Lily Lake Diatoms 21 Diatoma vulgare var. breve Grunow: Valve 28-32 [xm long by 12 jum wide; costae 7-9 in 10 jum; striae 1-2 between costae (Pa- trick and Reinier 1966:110). Frustnles were observed in samples from 150 cm only. Meridion circulare (Grev.) Agardh: Valve 26-28 jLim long by 5-6 jum wide; costae 3-5 in 10 jum; striae 16-18 in 10 jiim (Patrick and Reimer 1966:113). One or two specimens were observed in modern plankton samples. Fragilaria brevistriata Grunow: Valve 20-23 jam long by 3-3.5 /xm wide; striae 14 in 10 jam (Patrick and Reimer 1966:128). Fnis- tules were scattered in samples from 240 to 15 cm. The maximum relative density of 2.8 percent was from 150 cm. Fragilaria brevistriata var, inflata (Pant.) Hustedt: Valve 9-15 jttm long by 4-5 jum wide; striae 10-14 in 10 jum (Patrick and Rei- mer 1966:129). pRistules were scattered in samples from 360 to 75 cm. Their maximum relative density of 9 percent was at 135 cm. Fragilaria constricta Ehrenberg: Valve 40 jum long by 8 jum wide; striae 18 in 10 jum (Patrick and Reimer 1966:122). Frustules were very rarely observed in the sample from 165 cm. Fragilaria constricta f. stricta (A. CI.) Hustedt: Valve 24-28 jum long by 11-13 jum wide; striae 16-18 in 10 jum (Patrick and Rei- mer 1966:123). Fmstules were rare and ob- served only from 150 cm. Fragilaria construens var. venter (Ehr.) Grunow: Valve 7-10 /xm long by 4-5 /xm wide; striae 14-16 in 10 /im (Patrick and Rei- mer 1966:126). It was abundant throughout the core except in modern plankton samples. The maximum relative density of 18.4 per- cent was from 240 cm. Fragilaria crotonensis Kitten: Valve 43-45 /xm long by 2-3 /im wide; striae 14 in 10 /xm (Patrick and Reimer 1966:121). Frustules were rarely observed in samples from 360 to 270 cm. It constituted a maximum of only 0.2 percent of the diatom population. Fragilaria intermedia Grunow: Valve 39 /xm long by 8 /im wide; striae 11-12 in 10 /xm (Hustedt 1930:139). A single fnistule was ob- served at 150 cm. Fragilaria lapponica Grunow: Valve 19-24 /xm long by 5-6 /xm wide; striae 9-10 in 10 /im (Patrick and Reimer 1966:130). It was observed from 360 to 75 cm. Fragilaria leptostauron (Ehr.) Hustedt: Valve measurements not obtained (Patrick and Reimer 1966:124). It was observed as a single specimen in a sample from 135 cm. Fragilaria virescens Ralfs: Valve 13-35 /xm long by 5-10 /irn wide; striae 14-22 in 10 /xm (Patrick and Reimer 1966:119). Frustules were abundant in samples from all levels ex- cept from 105 to 75 cm and in the modern plankton. The maximum percent relative density of 18.4 percent was observed in sam- ples from 195 cm. Synedra amphicephala Kiitzing: Valve 22 /im long by 3 /xm wide; striae 16 in 10 /xm (Hustedt 1930:156). A single frustule was ob- served at 270 cm. Synedra rumpens Kiitzing: Valve 38 /im long by 2 /im wide; striae 16-18 in 10 /xm (Hustedt 1930:156). A single frustule was ob- served at 270 cm. Synedra tenera W. Smith: Valve 46-47 /xm long by 3-3.5 /im wide; striae 22-24 in 10 /xm (Patrick and Reimer 1966:137). Frustules were scattered in samples from 180 to 75 cm. They constituted a maximum of 1.2 percent of the diatom population from 135 cm. Synedra ulna (Nitzsch) Ehrenberg: Valve 6 /tm wide; striae 10 in 10 /im (Patrick and Reimer 1966:148). No entire frustules were found, although fragments were observed in samples from 90 cm. Synedra species 1: Valve curved, 80-100 /im long, 9 /im wide; axial area distinct, lin- ear, hyaline; central area absent; striae 18 in 10 /xm, parallel throughout valve. Specific identification of this taxon was not possible and no entire specimens were observed. Fragments were scattered from 315 to 195 cm with very low abundance. Synedra species 2: Valve 26-35 /xm long by 2-5 /xm wide; striae 14-18 in 10 /xrn. Frus- tules were similar to S. capitata in shape but were smaller. They were scattered in samples from 240 to 195 cm, where they constituted a maximum of 0.2 percent of the total diatom population. EUNOTIACEAE Eunotia arcus Ehrenberg: Valve 12-35 /im long by 4-5 /tm wide; striae 13-14 in 10 /xm (Patrick and Riemer 1966:212). Frustules were observed in samples only from the 210 cm level. 22 Great Basin Naturalist Vol. 39, No. 1 Eunotia arcus var. uncinata (Ehr.) Gru- now: Valve 21 jum long by 5 jum wide; striae about 14 in 10 jum (Patrick and Reimer 1966:213). It was observed as a single speci- men in a sample from 270 cm. Eunotia curvata (Kiitz.) Largest.: Valve 30-32 jum long by 2-4jLim wide; striae 14-15 in 10 jum (Patrick and Reimer 1966:189). Frustules were scattered throughout the core. The maximum relative density of 6 percent was observed in modern samples. Eunotia exigua (Breb. ex Kutz.) Ra- benhorst: Valve 12-35 /xm long by 4-5 ju,m wide; striae 18-20 in 10 jum (Patrick and Rei- mer 1966:215). It was common throughout the core and was also observed in modern samples. Its maximum relative density of 18.4 percent was observed from 60 cm. Eunotia hexaglyphis Ehrenberg: Valve 35-45 jLim long by 8 jum wide; striae 16 in 10 jum (Patrick and Reimer 1966:203). It was ob- served from 225 to 195 cm only. Its max- imum relative density of 3.2 percent was from 195 cm. Eunotia incisa W. Smith ex Greg.: Valve 30-33 fxm long by 5-5.5 jum wide; striae about 19 in 10 jum (Patrick and Reimer 1966:208). It was observed in three distinct zones from 285 to 195 cm, 60 to 45 cm, and in modern samples. In addition, a single frus- tule was observed at 360 cm. The maximum relative density was 2.2 percent at 195 cm. Eunotia lapponica Grunow ex A. Cleve: Valve about 50 jum long by 6 jum wide; striae 20 in 10 jum (Patrick and Reimer 1966:192). Frustules were observed from 225 to 165 cm and from 120 to 15 cm. The maximum num- bers were observed from 30 cm, where they constituted 9.6 percent of total diatom popu- lation. Eunotia maior (W. Sm.) Rabenhorst: Valve 80-168jum long by 8-14 jum wide; striae 9-10 in 10 jum (Patrick and Reimer 1966:196). Specimens were rare in the sample from 180 cm. Eunotia naegelii Migula: Valve 57 jum long by 3 jum wide; striae 18 in 10 jum (Pa- trick and Reimer 1966:190). A single fnistule was observed in a sample from 60 cm. Eunotia nymanniana Grunow: Valve 22 /x long by 3 jum wide; striae 14 in 10 jum (Pa- trick and Reimer 1966:214). A single speci- men was observed in a sample from 150 cm. Eunotia pectinalis var. minor (Kiitz.) Ra- benhorst: Valve 30-32 jum long by 5-6 jum wide; striae 16-18 in 10 jum (Patrick and Rei- mer 1966:207). E. pectinalis var. minor was observed at 285 cm and from 60 to 45 cm. The maximum relative density of 2 percent was observed at 60 cm. Eunotia praerupta var. bidens (Ehr.) Gru- now: Valve 60 jum long by 12-13 jum wide; striae 12 in 10 jum (Patrick and Reimer 1966:194). Frustules were observed only at 210 cm and 60 cm, where they constituted 0.8 percent and 0.2 percent of the total dia- tom population respectively. Eunotia praerupta var. inflata Grunow: Valve 33-53 jum long by 10-16 jum wide; striae 7-9 in 10 jum (Patrick and Reimer 1966:194). Frustules were scattered in sam- ples from 210 to 15 cm. They constituted 4 percent of the flora at 30 cm. Eunotia septentrionalis Oestrup: One bro- ken specimen was observed at 135 cm (Pa- trick and Reimer 1966:212). Eunotia serra var. diadema (Ehr.) Patrick: Valve 30-45 jum long by 11-18 jum wide; striae 11-12 in 10 jum (Patrick and Reimer 1966:201). It was observed mainly from 270 to 180 cm. A single frustule was found from 75 cm. A maximum relative density of 3.4 percent was observed from 195 cm. Eunotia suecica A. Cleve: Valve 27 jum long by 9 jum wide; striae 14 in 10 jum. (Pa- trick and Reimer 1966:199). One specimen was observed from 210 cm. Eunotia valida Hustedt: Valve 57-70 jum long by 5-6 jum wide; striae 12-14 in 10 jum (Patrick and Reimer 1966:192). It was scat- tered in samples from 285 to 30 cm and was also in the modern plankton. A maximum rel- ative density of 5.2 percent was observed from 210 cm. Eunotia vanheurckii Patrick: Valve 35-38 jum long by 9-10 jum wide; striae 16-20 in 10 jum (Patrick and Reimer 1966:210). It was scattered in lower samples from 345 to 195 cm. A few frustules were observed from 60 cm. A maximum relative density of 1.6 per- cent was observed from 195 cm. Eunotia vanheurckii var. intermedia (Krasske ex Hust.) Patrick: Valve 15-16 jum long by 3-4 jum wide; striae 16 in 10 jum (Pa- trick and Reimer 1966:211). It was observed in samples from 285 to 270 cm. March 1979 Jatkar et al.: Lily Lake Diatoms 23 ACHNANTHACEAE Cocconeis pediculus Ehrenberg: Valves 25-26 jixm long by 17-21 jum wide; striae 19-20 in 10 fxm (Patrick and Reimer 1966:240). Frustules were observed from 360 cm and from 150 to 135 cm. At 150 cm it constituted 2.2 percent of total diatom popu- lation. Cocconeis placentula var. eugylpta (Ehr.) Cleve: Valve 13-26 jum long by 8-20 jum wide; striae 19-24 in 10 jum (Patrick and Rei- mer 1966:241). It was observed from 180 to 90 cm. It had a maximum relative density of 4.8 percent from 150 cm. Two frustules were additionally observed in the modern plankton sample. Achnanthes biasolettiana Kiitzing: Valve 12-12.5 jum long by 4 jxm wide; striae 18 in 10 jLim (Hustedt 1930:199). A few frustules were observed at 120 cm. Achnanthes conspicua A. Mayer: Valve 7-8 jLtm long by 4 jum wide; striae 18 in 10 jum on raphe valve (Hustedt 1930:202). Frus- tules were observed only from 270 cm, where they constituted 1.4 percent of total diatom population. Achnanthes delicatula Kiitzing: Valve 16-17 jum long by 5-5.5 jum wide; striae 15-17 in 10 jum on raphe valve (Hustedt 1930:202). Frustules were observed only from 240 cm, where they constituted 0.7 percent of the total diatom population. Achnanthes linearis f. curta H. L. Smith: Valve about 10 jum long by 2.5-3.5 jum wide; striae about 25 in 10 jum (Patrick and Reimer 1966:252). Fmstules were scattered in sam- ples from 360 to 225 cm, 150 cm, 90 cm, and the extant plankton. The maximum relative density of 7.7 percent for this taxon was ob- served from 255 and 240 cm. Achnanthes linearis var. pusilla Grunow: Valve 15-16 jum long by 3.5-5 jum wide; striae 22-24 in 10 jum (Patrick and Reimer 1966:252). It was observed only from 360 to 225 cm. The maximum relative density of 9.1 percent was observed at 315 cm. Achnanthes minutissima Kiitzing: Valve 9-11 jum long by 2.5 jum wide; striae very fine (Hustedt 1930:198). Frustules were ob- served at 120 cm. Achnanthes minutissima var. cryptoce- phala Grunow: Valve 13-18 jum long by 2-3.5 jum wide; striae very fine. (Hustedt 1930:198). Frustules were observed only from 270 to 225 cm. The maximum relative den- sity was 7.9 percent at 240 cm. Naviculaceae Mastogloia elliptica Agardh: Valve 25-38 jum long by 10 jum wide; striae about 16 in 10 jum (Hustedt 1930:217). M. elliptica frustules were rare in samples from 60 cm. Frustulia rhomboides (Ehr.) de Toni: Valve 120-125 jum long by 20-22 jum wide; striae 20-25 in 10 jum (Patrick and Reimer, 1966:306). F. rhomboides frustules were scat- tered in the deeper section of the core from 360 cm to 165 cm. This taxon constituted 3.6 percent of the total diatom population from 300 cm. Frustulia rhomboides var. capita (A. May- er) Patrick: Value 39-45 jum long by 10-11 jum wide; striae about 30 in 10 jum (Patrick and Reimer 1966:307). Frustules of this taxon were scattered throughout the core. A max- imum relative density of 16.2 percent oc- curred in the surface sample. Frustulia rhomboides var. crassinervia (Breb. ex W. Sm.) Ross: Valve 45-46 jum long by 10-10.5 jum wide; striae very fine, unresol- vable (Patrick and Reimer 1966:307). It was observed in samples from 180 to 15 cm and also in the surface sample. A maximum fre- quency of 18.8 percent was from 90 cm. Caloneis limosa (Kiitz.) Patrick: Valve 55-70 jum long by 10-12 jum wide; striae 16-20 in 10 jum (Patrick and Reimer 1966:587). One or two specimens of this tax- on were observed from 360 cm. Neidum bisulcatum (Lagerst.) Clave: Valve 50-70 jum long by 9-12 jum wide; striae 26-30 in 10 jum (Patrick and Reimer 1966:397). A few specimens of N. bisulcatum were observed from 360 cm. Neidium iridis (Ehr.) Cleve: Valve 50-80 jum long by 14-20 jum wide; striae about 20 in 10 jum (Patrick and Reimer 1966:386). It was only in samples from the lower section of the core (360 to 225 cm). A maximum rela- tive density of 2 percent was observed in samples from 360 cm. Neidium iridis var. amphigomphus (Ehr.) A. Mayer: Valve 45-122 jum long by 16-25 24 Great Basin Naturalist Vol. 39, No. 1 jLim wide; striae 16-18 in 10 jum (Patrick and Reimer 1966:387). It was observed mainly from the deeper section of the core (345 to 210 cm). A maximum relative density of 3.2 percent was observed from 240 cm. A few frustules were from 30 cm, 15 cm, and the modern plankton sample. Neidium iridis var. ampliatum (Ehr.) Cleve: valve 35-85 jum long by 12-20 /xm wide; striae about 20 in 10 jum (Patrick and Reimer 1966:388). It was observed from 330 and 315 cm where it constituted about 3.5 percent of the diatom population. A few frus- tules were from 255 cm and in the extant plankton. Diploneis elliptica (Kiitz.) Cleve: Valve 37 jLim long by 25 jum wide; costae 8-9 in 10 jitm; alveoli in single row between costae (Patrick and Reimer 1966:414). It was observed only from 165 cm and 105 cm, where it occurred as single specimens. Diploneis interrupta (Kiitz.) Cleve: Valve 55 jum long by 8-9 ju,m wide; costae 8 in 10 jum (Patrick and Reimer 1966:416). A single entire frustule was observed from 75 cm. Diploneis smithii var. dilatata (M. Perag.) Beyer: Valve 36-37 [xm long by 18-18.5 jum wide; costae 10 in 10 jum; double row of al- veoli between costae (Patrick and Reimer 1966:411). Frustules were observed only from 180 cm and 135 cm. The maximum relative density of 1.2 percent was observed from 135 cm. Stauroneis anceps Ehrenberg: Valve about 70 jum long by about 14 [xm wide; striae 18 in 10 jLim (Patrick and Reimer 1966:361). It was observed mostly from 360 to 135 cm. The maximum frequency of 1.6 percent was ob- served from 345 cm. Stauroneis anceps f. gracilis Rabenhorst: Valve to 67 ]u,m long by up to 14 /xm wide; striae very fine (Patrick and Reimer 1966:361). It was observed only in the deeper section of the core (from 360 to 240 cm). It constituted 0.9 percent of the diatom popu- lation from 315 cm and 240 cm. Stauroneis kriegeri Patrick: Valve 12-23 jLim long by 3 jum wide; striae about 26-28 in 10 jLim (Patrick and Reimer 1966:362). Very few specimens were observed only from the upper core. Stauroneis phoenicenteron (Nitz.) Ehren- berg: Valve 155-157 jum long by 25-26 jum wide; striae 14-15 in 10 jum (Patrick and Rei- mer 1966:359). It was observed from 330 to 285 cm and from 210 to 195 cm. This diatom constituted 1.2 percent of the total popu- lation from 285 cm. A single frustule was in the extant plankton. Anomoeoneis costata (Kiitz.) Hustedt: Valve 75 jum long by 25 jum wide; striae 16 in 10 jum (Patrick and Reimer 1966:376). Only one or two specimens were observed from 90 cm. Anomoeoneis serians var. acuta Hustedt: Valve 29-40 jum long by 7.5-8 jum wide; striae 26-28 in 10 jum (Patrick and Reimer 1966:378). This taxon was observed in extant samples only where it constituted 4.6 percent of the diatom population. Anomoeoneis serians var. brachysira (Breb. ex Kutz.) Hustedt: Valve 15-32 jum long by 5-6.5 jum wide; striae 24-30 in 10 jum (Patrick and Reimer 1966:379). Frustules were abundant throughout the core. A max- imum frequency of 16.4 percent was ob- served from 30 cm. Anomoeoneis vitrea (Grun.) Ross: Valve 25-32 jum long by 5-6 jum wide; striae 30 or finer in 10 jum (Patrick and Reimer 1966:380). Frustules were very common in deeper parts of the core (from 360 to 225 cm). This taxon constituted 3.2 percent of the flora from 225 and 240 cm. Navicula aikenensis Patrick: Valve 18-25 jum long by 5-6 /xm wide; striae about 16 in 10 /im (Patrick and Reimer 1966:473). It was observed only from 360 to 225 cm. A max- imum relative density of 1.2 percent was ob- served from 285 cm. Navicula angusta Grunow: Valve 55-62 /im long by 5-7 /im wide; striae 12-14 in 10 /im (Patrick and Reimer 1966:514). It was ob- served only from 285 cm and from 240 cm to 225 cm. A maximum relative density of 1.4 percent was observed from 285 cm. Navicula hofleri Cholnoky: Valve 31-32 /im long by 8-8.5 /im wide; striae very fine (Hustedt 1961-1966:97). Frustules were scat- tered throughout the core. A maximum rela- tive density of 9.6 percent was observed from 15 cm. Navicula mutica Kiitzing: Valve 13-15 /xm long by 5-6 /im wide; striae about 20 in 10 /xm (Patrick and Reimer 1966:454). It was ob- served from 195 to 45 cm. A maximum rela- March 1979 Jatkar et al.: Lily Lake Diatoms 25 tive density of 7.2 percent was observed from 135 cm. Navicula mutica var. cohnii (Hilse) Gru- now: Valve 14-15 jum long by 6-7 jum wide; striae 16-18 in 10 jum (Patrick and Reimer 1966:454). It was observed rarely at 285 cm. Navicular mutica var. undulata (Hilse) Grunow: Valve 9-16 jum long by 5-7 jum wide; striae 18-20 in 10 jum (Patrick and Rei- mer 1966:456). It was observed only at 240 cm and from 105 to 90 cm. Single fmstules were observed at each level. Navicula pelliculosa (Breb.) Hilse: Valve 6 jum long by 4 jum wide; striae very fine (Hus- tedt 1930:287). One specimen was observed at 105 cm. Navicula pseudoscutiformis Hustedt: Valve 12-13 ju,m long by 10 /xm wide; striae 20-22 in 10 jum (Patrick and Reimer 1966:451). It was observed from 285 to 240 cm. A maximum relative density of 2 percent was observed from 255 cm. Navicula pupula var. rectangularis (Greg.) Grunow: Valve 25-37 jum long by 8-9 ]um wide; striae 20 in 10 jum (Patrick and Reimer 1966:497). Frustules were scattered from 360 to 90 cm and a few were observed from sur- face samples. A maximum relative density of 19.4 percent was observed from 345 cm. Navicula radiosa Kiitzing: Valve 70-83 jLtm long by 10-12 jum wide; striae 12-13 in 10 jum (Patrick and Reimer 1966:509). It was observed mostly in deeper samples (360 to 225 cm). A few frustules were scattered in some of the upper and the extant samples. A maximum frequency of 5.6 percent was ob- served from 270 cm. Navicula subtilissima Cleve: Valve 20-28 jum long by 4-6 jum wide; striae very fine (Patrick and Reimer 1966:483). Frustules were scattered throughout the core but pri- marily in the upper strata. A maximum rela- tive density of 21 percent was observed from 150 cm. Navicula longirostris Hustedt: Valve 18-19 jum long by 3-3.5 jum wide; striae very fine (Hustedt 1930:285). Frustules occurred only from 315 to 240 cm. A maximum rela- tive density of 2.5 percent was observed at 315 and 285 cm. Pinnularia abaujensis (Pant.) Ross: Valve 60 jum long by 12 jum wide; striae 13-14 in 10 jum (Patrick and Reimer 1966:612). Frus- tules were observed rarely from 285-270 cm, from 210 cm, and from 90 cm. A maximum relative density of 0.6 percent was observed from 90 cm. Pinnularia abaujensis var. rostrata (Patr.) Patrick: Valve 60-62 jum long by 11-12 jum wide; striae to 14 in 10 jum (Patrick and Rei- mer 1966:614). Only one or two frustules were observed from 270 cm. Pinnularia biceps Greg.: Valve 39-71 jum long by 8-10 jum wide; striae 10-15 in 10 jum (Patrick and Reimer 1966:599). It was pres- ent throughout the core. A maximum relative density of 20.4 percent was observed from 330 cm. Pinnularia borealis Ehrenberg: Valve 32-44 jum long by 8.5-10 /xm wide; striae 5-6 in 10 jum (Patrick and Reimer 1966:618). Frustules were observed from 210 cm and from 150 to 45 cm. A maximum relative den- sity of 2.4 percent was observed from 135 cm. Pinnularia braunii (Grun.) Cleve: Valve 35-37 jum long by 8-9 jum wide; striae 12 in 10 jum (Patrick and Reimer 1966:594). It oc- curred only from 360 cm and 345 cm. A max- imum relative density of 1.7 percent was ob- served from 360 cm. Pinnularia brebissonii (Kiitz.) Rabenhorst: Valve 42-75 jum long by 7-11 jum wide; striae 8-14 in 10 jum (Patrick and Reimer 1966:614). A few specimens of P. brebissonii were observed at 360 cm and 255 cm. Pinnularia dactylus Ehrenberg: Valve 230-235 jum long by 28-30 jum wide; striae 4-5 in 10 jum (Patrick and Reimer 1966:632). Single frustules were observed from 285 cm to 255 cm only. Pinnularia divergens W. Smith: Valve 65 jum long by 15 jum wide; striae 12 in 10 jum (Hustedt 1930:323). It was observed at 345 cm. Pinnularia gibba Ehrenberg: Valve 60-100 jum long by 10-12 jum wide; striae 8-12 in 10 jum (Hustedt 1930:327). Frustules were observed from 360 to 210 cm. A max- imum relative density of 3.2 percent was ob- served from 285 cm. Pinnularia mesolepta f. angusta Cleve: Valve 26-60 jum long by about 5-7.5 jum wide; striae 12-20 in 10 jum (Hustedt 1930:319). A few frustules were observed at 360 cm. 26 Great Basin Naturalist Vol. 39, No. 1 Pinnularia microstauron (Ehr.) Cleve: Valve 52-63 ]um long by 10-12 jum wide; striae 9-13 in 10 jum (Patrick and Reimer 1966:597). It was observed in all strata. At 45 cm these frustules constituted 46 percent of the total diatom population. Pinnularia molaris Grunow: Valve 30 ]u,m long by 7 ju,m wide; striae 14 in 10 jum (Hus- tedt 1930:316). A few frustules were ob- served in the extant plankton sample. Pinnularia nobilis (Ehr.) Ehrenberg: Valve 280-282 /xm long by about 30 jum wide; striae to 7 in 10 jum (Patrick and Rei- mer 1966:638). Single frustules of P. nobilis were observed from 165 cm and 120 cm. Pinnularia platycephala (Ehr.) Cleve: Valve 77 jum long by 18 jum wide; striae 10 in 10 jum (Hustedt 1930:324). A single frustule was observed at 360 cm. Pinnularia subsolaris (Grun.) Cleve: Valve 75 jum long by 15 jum wide; striae 10 in 10 jLim (Hustedt 1930:322). Frustules were ob- served only from 360 and 345 cm. A max- imum frequency of 0.5 percent was observed from 360 cm. Pinnularia viridis (Nitzsch) Ehrenberg: Valve 120-175 jum long by 21-30 jum wide; striae 5-9 in 10 ju,m (Patrick and Reimer 1966:639). It was scattered in the deeper sec- tion of the core (from 360 to 210 cm only). A maximum relative density of 7.4 percent was observed from 210 cm. Pinnularia viridis var. commutata (Grun.) Cleve: Valve 42-90 fim long by 9-17 fxm wide; striae 9-14 in 10 jxm (Patrick and Rei- mer 1966:640). It was observed at 360 cm. Pinnularia viridis var. minor Cleve: Valve 85-100 jum long by about 18 jum wide; striae 8-9 in 10 nm (Patrick and Reimer 1966:641). It was observed from 135 to 15 cm. A max- imum relative density of 12.2 percent was observed from 75 cm. Pinnularia species 1: Valve 25-35 jum long by 9-10 jum wide; striae 12-13 in 10 jum. It was scattered from 360 to 180 cm. At 360 cm maximum relative density of 6.7 percent was observed. Pinnularia species 2: Valve 40 jum long by 10 jum wide; striae 12 in 10 jum; isolated punctum in central area. A few frustules were observed at 270 cm. Cymbellaceae Cymbella amphicephala Naeg. ex Kiitzing: Valve 30-32 /xm long by 8-10 jum wide; striae 14-16 in 10 jum (Patrick and Rei- mer 1975:33). Frustules were observed only in deeper strata (from 360 cm to 195 cm). A maximum frequency of 1.7 percent was ob- served from 360 cm. Cymbella angustata (W. Sm.) Cleve: Valve 35-43 jum long by 8-9 jum wide; striae 16-20 in 10 jum (Patrick and Reimer 1975:22). It was observed from 345 cm to 210 cm. A few faistules were also observed from 165 cm and 30 cm. A maximum relative density of 3.1 percent was observed from 315 cm. Cymbella cuspidata Kiitzing: Valve 80-90 jum long by 22 jum wide; striae 13-14 in 10 jum (Patrick and Reimer 1975:39). Frustules were observed only at 285 and 270 cm. Their relative density was 0.5 percent. Cymbella hebridica Grun. ex Cleve: Valve 28-35 ]um long by 7-8 jum wide; striae 11-16 in 10 jum (Patrick and Reimer 1975:30). It was observed from 330 to 15 cm and in ex- tant samples. A maximum relative density of 14 percent was observed from 165 cm. Cymbella heteropleura var. subrostrata Cleve: Valve 175-177 jum long by 45-47 jum wide; striae 11-12 in 10 jum (Patrick and Rei- mer 1975:38). It was observed from 315 to 255 cm and at 195 cm. A maximum relative density of 0.5 percent was observed from 285 cm. Cymbella lunata W. Smith: Valve 35-55 jum long by 5-8 jum wide; striae 10-14 in 10 fxm (Patrick and Reimer 1975:46). Frustules were scattered from 360 to 60 cm and were also observed in extant samples. A maximum relative density of 10.2 percent was observed from 345 cm. Cymbella minuta Hilse ex Rabenhorst: Valve 18-19 jum long by 5-7 jum wide; striae 12-14, becoming 18-19 in 10 jum (Patrick and Reimer 1975:47). It was observed from 360 to 255 cm. A maximum relative density of 9.1 percent was observed from 360 cm. Cymbella minuta f. latens (Krasske) Rei- mer: Valve 23-24 jum long by 6-7 jum wide; striae 12 in 10 jum (Patrick and Reimer 1975:49). It was scattered from 330 to 105 cm and was also observed in extant samples. March 1979 Jatkar et al.: Lily Lake Diatoms 27 The maximum relative density of 2.4 percent was observed from 180 cm. Cymbella naviculiformis Auersw. ex Heib.: Valve 40-42 jum long by 10-11 jitm wide; striae 13-14 in 10 jum (Patrick and Rei- mer 1975:31). It was observed from 330 and 315 cm, 180 cm, and from 135 to 75 cm. The maximum relative density of 6 percent was observed from 330 cm and 135 cm. Cymbella perpusilla A. Cleve: Valve 16 jLim long by 4 jum wide; striae to 16 in 10 jum (Hustedt 1930:361). Frustules were observed only from 300 cm to 225 cm and from 150 cm. The maximum relative density of 6.2 percent was observed from 150 cm. Cymbella sinuata Gregory: Valve mea- surements not recorded (Patrick and Reimer 1975:51). One specimen was observed from 195 cm. Cymbella turgida (Greg.) Cleve: Valve 52 jLim long by 12 jitm wide; striae 10 in 10 jum (Hustedt 1930:358). One frustule was ob- served from 315 cm. Amphora ovalis (Kiitz.) Kiitzing: Valve 33-40 jLtm long by 10-12 jum wide; striae 12 in 10 jum (Patrick and Reimer 1975:68). One frustule was observed at each of four levels and also in the extant plankton. GOMPHONEMACEAE Gomphonema acuminatum Ehrenberg: Valve 35-40 jum long by 7-8 ju.m wide; striae 12 in 10 JLtm (Patrick and Reimer 1975:112). One frustule was observed at 270 cm. Gomphonema affine Kiitzing: Valve 30 jum long by 7 jum wide; striae 14 in 10 jum (Patrick and Reimer 1975:133). Only one or two frustules were observed at 240 cm. Gomphonema angustatum var. intermedia Grunow: Valve 27 jum long by 8 jum wide; striae 12 in 10 jum (Patrick and Reimer 1975:126). One specimen was observed from 135 cm. Gomphonema apicatum Ehrenberg: Valve 35-36 jum long by 8 pun wide; striae 12-13 in 10 jum (Patrick and Reimer 1975:110). Frus- tules were observed only from 210 cm and 180 cm where their relative density was 0.4 percent and 0.2 percent respectively. Gomphonema gracile Ehrenberg em. V. H.: Valve 40-42 jum long by 6.5-7 jum wide; striae 16 in 10 jum (Patrick and Reimer 1975:131). Frustules were scattered from 345 cm to 165 cm. A maximum relative density of 0.7 percent was observed from 285 cm. Gomphonema grunowii Patrick: Valve 38 jum long by 7 jum wide; striae 12 in 10 jum (Patrick and Reimer 1975:131). It was ob- served from 330 cm, 300 to 285 cm, and 210 cm. A maximum relative density of 0.5 per- cent was ob.se rved from 330 cm. Gomphonema olivaceum (Lyngb.) Kiitzing: Valve 23-25 jum long by 6.5-7 jum wide; striae 12 in 10 jum (Patrick and Reimer 1975:139). Frustules were observed from 360 cm, 315 cm, and 180 to 135 cm. The max- imum relative density of 6.4 percent was ob- served from 150 cm. Gomphonema parvulum (Kiitzing): Valve 29 jum long by 6 jum wide; striae 12 in 10 jum (Patrick and Reimer 1975:122). Only one or two frustules were observed at 270 cm. Gomphonema subtile Ehrenberg: Valve 43-45 ]um long by 5 jum wide; striae 16 in 10 ]um (Patrick and Reimer 1975:117). A few specimens were observed from 270 cm. Gomphonema truncatum var. capitatum (Ehr.) Patrick: Valve 25-50 jum long by 10-15 jum wide; striae about 13 in 10 jum (Pa- trick and Reimer 1975:119). Frustules were scattered between 330 and 195 cm and were also found from 135 cm. The maximum rela- tive density of 1.2 percent was observed from 195 cm. One frustule was in the extant plank- ton. Epithemiaceae Denticula elegans f. valida Pedicino: Valve 35-55 jum long by 5-10 jum wide; cos- tae 3-4 in 10 jum; striae 5-6 between costae (Patrick and Reimer 1975:171). One or two frustules were observed from 180, 150, and 120 cm. Rhopalodia gibba (Ehr.) O. Miiller: Valve 120-155 jum long by 7-8.5 jum wide; costae 7 in 10 jum; striae 1-2 between costae (Patrick and Reimer 1975:189). One frustule was ob- served from 135 cm and three were observed from 105 cm and 90 cm. Rhopalodia gibberula var. protracta Gru- now: Valve 40-60 jum long by 8-11 jum wide; costae 3-4 in 10 jum; striae 14-16 in 10 jum (Hustedt 1930:391). It was rare from 150, 135, 90, and 75 cm. 28 Great Basin Naturalist Vol. 39, No. 1 Epithemia sorex Kiitzing: Valve 35-37 jum long by 9-9.5 jum wide; costae 5 in 10 jum; 3-4 rows of alveoli between costae (Patrick and Reimer 1975:188). It was observed from 135 cm and 105 to 75 cm. At 105 cm the maximum relative density of 1 percent was observed. Epithemia turgida (Ehr.) Kiitzing: Valve 140-145 jum long by 18-20 ju,m wide; costae 4 in 10 jum; two rows of alveoli between costae (Patrick and Reimer 1975:182). Frustules were observed from 150 to 75 cm. A max- imum relative density of 1.2 percent was ob- served from 120 cm. Epithemia turgida var. westermannii (Ehr.) Grunow: Valve 70-72 jtim long by 14 jLim wide; costae 4 in 10 /xm; two rows of al- veoli between costae (Patrick and Reimer 1975:184). One frustule was observed from 165 cm. Nitzschiaceae Nitzschia dissipata (Kiitz.) Grunow: Valve 24 jum long by 3-4 jum wide; keel punctae 8-10 in 10 jLim; striae very fine (Hustedt 1930:412). A few frustules were observed at 120 cm. Nitzschia fonticola Grunow: Valve 9-16 jum long by 3-3.5 /xm wide; keel punctae 12-14 in 10 ]um; striae very fine (Hustedt 1930:415). Frustules were observed from 345 to 210 cm. A few frustules were scattered at 165, 90, and 30 cm. The maximum relative density of 6 percent was observed from 285 cm. Nitzschia frustulum (Kiitz.) Grunow: Valve 20-22 /xm long by 3-4 jum wide; keel punctae 11-12 in 10 jum; striae 20 in 10 ju,m (Hustedt 1930:415). It was only in deeper strata from 360 to 225 cm. The maximum rel- ative density of 7.3 percent was obtained from 240 cm. Nitzschia palea (Kiitz.) W. Smith: Valve 30-32 jLim long by 2.5-3 /xm wide; keel punc- tae 12 in 10 jLxm; striae very fine (Hustedt 1930:416). Frustules were scattered from 345 to 90 cm, at 15 cm and in extant samples. The maximum relative density of 5.2 percent was observed from 180 and 15 cm. Nitzschia species: Valve linear-lanceolate, 65-67 jLim long by 7-7.5 /xm wide; keel punc- tae 8 in 10 jLim; striae 20 in 10 jum. Specific identification of this taxon was not possible. It resembles N. thennalis in general shape but differs in shape of the poles. It also resembles N. capitellata in general shape but the num- ber of striae is different. It resembles N. an- gustata in shape but the striae in N. angus- tata are much coarser. One frustule was found from 360 cm, 5 from 150 cm, and 8 from 135 cm. Hantzschia amphioxys f. capitata O. Miiller: Valve 30-52 jum long by 7-10 jum wide; keel punctae 8-10 in 10 jum; striae about 20 in 10 ium (Hustedt 1930:394). Frus- tules were observed from 225 to 60 cm. At 135 cm they constituted 16.2 percent of the diatom population. SURIRELLACEAE Cymatopleura solea (Breb.) W. Smith: Valve 65-70 jum long by 10-15 /xm wide; cos- tae 4-6 in 10 /tm; striae not clearly observed (Hustedt 1930:425). Frustules were observed at 285 cm. Surirella ovalis Brebisson: Valve measure- ments not recorded (Hustedt 1930:441). One frustule was observed at 120 cm. Surirella robusta Ehrenberg: Valve 120-145 /xm long by 40-65 /xm wide; costae 1-2 in 10 /xm; striae 8-12 in 10 /tm (Hustedt 1930:437). A few frustules were observed at 360 cm. Distribution and Abundance The occurrence and abundance of diatoms in 27 samples taken from the core were de- termined. The highest counts were for non- planktonic species, belonging mainly to the genera Cymbelki, Pinnularia, Navicula, Gom- phonema, and Eunotia. Planktonic forms en- countered belonged mainly to the genera Cyclotella, Melosira, Fragilaria, Steph- anodiscus and Tabellaria. The abundance of all taxa whose frequency was four or more in any sample is shown in Figure 2. This figure is prepared with plank- tonic taxa on the right and periphytic taxa on the left. The three most abundant species by far in this study were Navicula subtilissima, Pinnularia microstauron, and Tabellaria fen- estrata. Anomoeoneis serians var. brachysira, Cymbella hebridica, Cymbelki lunata, Fragi- March 1979 Jatkar et al.: Lily Lake Diatoms 29 ruitulio rhomboid** Frufttulio rhomboidvt var. craitinsrvi Oomphonomo loncoolotu Bviculo hbfflori viculo pupulo vor. roctongularic Naviculo longiroilrii Noidium iridif vor. omphigomphuf Noidium iridii var. ampliatum Nitiichio fonlicola Nitiachio trullulum Nitstchio fuiilormir Nititchio pal«a Nitiichia apccici Pinnuloria bi<*pt Pinnulorio beroolU Pinnuloria viridii Pinnuloria gibbo oncopi vor. gracilii .n*i. ph C 2 Fig. 2. Percent frequency of individual diatom taxa according to sediment depth. Euplankters are listed at the top of the figure. Scale of frequency width is shown on the bottom right. 30 Great Basin Naturalist Vol. 39, No. 1 Cyclotcllo Kullingiono br«vi«triata vor. br«viitriata liloria br*viitriata vor. inflota agilori« vir*ft<*n« Mcloiira dickici M»losira diitont Tab*llaria <*n«itrata Tsbdlsria flocculoia Achnonthss lin«arii f. curto Achnonth** lin*arit vor. puiilia AchnanlhM minutiiiimo vor. cryptocaphala Achnanth** contpicuo A 5 ^°- o Q. (0 i.oj I I I I r 300 I I I I I I I I I I I 120 30 0 Depth Fig. 4. Shannon- Weaver diversity indices for diatom populations in relation to sediment depth. r = 0.754, which significant at the 0.01 level. 34 Great Basin Naturalist Vol. 39, No. 1 %> Similarity r 1 l| H 1 — — 1 1 r >i . VI Fig. 5. Phenogram (cluster diagram) demonstrating similarity between samples from different sediment depths. Sample numbers are shown within subgroups and correspond to depth, with sample 2 being the deepest (oldest) and samples 26 and 27 being surface samples. March 1979 Jatkar et al.: Lily Lake Diatoms 35 riod was not as homogenous as the others, and depositional environments apparently fluctuated and eventually disrupted the de- velopment of acidic conditions. This distur- bance intermpted the development of strictly dytrophic conditions and apparently shifted the system toward a new period of circum- neutral mesotrophic (?) conditions. Recent conditions in the lake were again less stable but appear to be returning to a more acidic, dystrophic state. This could explain why our surface samples clustered with those of deep- er sediments (in subgroup B). Prevalent Species and Cluster Analysis A list of species encountered in this study and identified as prevalents is given in Table 3. Prevalent species were selected based upon their occurrence in 30 percent or more of the samples. These species were subjected to niche breadth and niche overlap analyses (Colwell and Fatuyma 1971; Table 3). Spe- cies were then clustered based upon niche overlap values (Cody 1974). This analysis represents an attempt to group diatom spe- cies according to preference for similar envi- ronmental and/or habitat conditions. Three Table 2. Clusters (subgroups) as delineated on Figure 5. The prominent species responsible for the uniqueness of subgroups are shown. Species Cluster subgroups (Fig. 5) Pinnularia biceps Fragilaria construens var. venter Achnanthes linearis f. curta Anomoeoneis vitrea Cymbella lunata Navictila pupula var. rectangularis Cymbella minuta Nitzschia fonticola Fragilaria virescens Achnanthes linearis var. pusilla Pinnularia species 1 Navictila radiosa Nitzschia frustulum Cymbella angustata Pinnularia species 2 Achnanthes minutissima var. cryptocephala Neidium iridis var. amphigomphus Tabellaria fenestrata Navicula subtilissima Frustulia rhomboides var. capitata Eunotia curvata Anomoeoneis serians var. acuta Eunotia serra var. didyma Arumioeoneis serians var. brachysira Pinnularia microstauron Cymbella hebredica Eunotia lapponica Tabellaria flocculosa Eunotia valida Eunotia exigua Pinnularia viridis Pinnularia viridis var. minor Navicula mutica Frustulia rhomboides Nitzschia palea Hantzschia amphioxys f. capitata Pinnularia borealis Fragilaria brevistriata var. inflata A B C D 11.17 1.58 0.57 0.32 8.81 2.08 0.86 0.10 3.21 0.68 0.60 0.0 2.97 0.10 0.0 0.0 6.76 4.12 0.20 0.04 3.88 0.24 0.34 0.04 2.21 0.0 0.0 0.0 3.37 0.14 0.11 0.24 8.88 8.74 0.66 1.36 2.12 0.22 0.0 0.0 1.19 0.08 0.0 0.0 3.51 2.18 0.83 0.0 3.67 0.18 0.0 0.0 1.42 0.04 0.0 0.16 1.60 0.26 0.0 0.12 1.98 0.14 0.0 0.0 1.24 0.18 0.03 0.08 4.66 17.86 1.51 8.48 1.02 8.20 10.97 0.0 0.89 4.66 2.34 0.40 0.09 2.46 0.31 0.0 0.0 1.76 0.0 0.0 0.22 1.22 0.03 0.36 4.43 4.96 2.80 7.48 1.28 0.90 4.54 25.84 1.73 3.42 5.77 7.76 0.0 1.20 1.14 6.28 0.0 2.48 0.0 1.76 0.06 0.60 0.23 2.56 0.13 1.02 1.60 9.80 0.30 0.04 0.0 1.48 0.0 0.0 5.49 3.48 0.02 0.20 3.46 0.40 0.02 0.56 7.63 0.80 1.30 2.14 2.37 0.0 0.0 0.36 7.29 0.44 0.0 0.0 1.29 0.44 0.0 0.20 2.8 0.0 'Figures are mean importance values as determined by multiplying presence times average relative frequency (Warner and Harper 1972). 36 Great Basin Naturalist Vol. 39, No. 1 Table 3. List of 45 species occurring in 30 percent or more of the samples. Relative niche breadth and niche overlap values are reported for all species. Relative Average Species Niche Niche Name Breadth Overlap Achnanthes linearis var. pusilla 53.701 31.245 Achnanthes linearis f. curta 68.383 30.171 Anomoeoneis serians var. brachysira 89.400 49.244 Anomoeoneis vitrea 66.324 38.018 Cyclotella kiitzingiana 71.928 28.309 Cymbella amphicephala 70.015 37.349 Cymbella angustata 66.395 38.523 Cymbella hebridica 86.469 44.138 Cymbella lunata 83.010 47.525 Cymbella mimita f. latens 72.314 39.631 Cymbella naviculiformis 55.657 28.322 Eunotia curvata 55.8,33 22.718 Eunotia exigua 73.128 28.090 Eunotia incisa 69.010 .33.792 Eunotia lapponica 74.453 29.151 Eunotia praerupta var. inflata 58.251 23.606 Eunotia valida 73.148 30.748 Eunotia vanheurcki 71.241 30.136 Fragillaria brevistriata 68.657 29.118 Fragillaria brevistriata var. inflata 64.588 19.859 Fragilaria construens var. venter 78.679 45.792 Fragilaria virescens 82.047 46.837 Frustulia rhomboides 73.972 40.406 Frustulia rhomboides var. capitata 73.770 36.5.34 Frustulia rhomboides var. crassinervia 67.992 28.289 Gomphonema truncatum var. capitatum 69.813 38.090 Hantzschia amphioxys f. capitata 64.866 28.329 Melosira granulata 67.492 29.150 Navicula heufleri 58..306 20.246 Navicula mutica 67.445 29.306 Navicula pupula var. rectangularis 62.431 33.632 Navicula radiosa 81.112 44.717 Navicula subtilissima 75.065 33.805 Neidium iridis var. amphigomphus 63.813 35.999 Nitzschia fonticola 71.934 41.987 Nitzschia frustulum 65.923 38.151 Nitzschia palea 81.747 43.753 Pinnularia biceps 75.766 44.059 Pinnularia borealis 68.856 29.436 Pinnularia microstauron 70.695 32.405 Pinnularia viridis var. minor 70.814 29.710 Pinnularia gibha 70.770 41.579 Pinnularia brehissonii 63.346 33.225 Tabellaria fenestrata 87.658 46.379 Tabellaria flocculosa 66.297 24.348 major clusters are evident from this analysis. Most species in the first cluster (group I, Fig. 6) flourished during the earliest period of sedimentation (from sample 2 at 360 cm to sample 12 at 210 cm). The majority of spe- cies in the cluster for which ecological pref- erences are known are generally considered to be pH indifferent or alkaliphilous, sap- roxenous to mesosaprobic and to prefer oli- gotrophic to eutrophic conditions. The spe- cies in this cluster correspond well with the species prominent in subgroup A (Table 2) of the sample cluster (Fig. 5). This is further evi- dence for the nondystrophic nature of Lily Lake early in its depositional history. The species in group II (Fig. 6) correspond well with the species which delineate sub- group C of Figure 5. As previously noted, subgroup C represents a time discreet unit of deposition which followed a short period of dystrophy and disturbance. The species for which ecological preferences are known in group II (Fig. 6) are indifferent to pH or pre- fer alkaline waters, are oligosaprobic to me- sosaprobic, and occur in mesotrophic to eu- trophic waters. These results show more strongly than sample clustering the fact of a second oligo-mesotrophic period in the lake history. This period corresponds approx- imately with deposition levels from 150 cm to 60 cm. Group III (Fig. 6) includes species from both subgroups B and D of Table 2 and Fig- ure 5. The species of group III with known ecological preferences are indicative of acid- ic waters and often dystrophic or bog habi- tats. This group is compo.sed of species found in midcore and surface sediments. The fact that these species overlap in their niche pref- erences even though widely separated depo- sitionally gives additional strength to our hy- pothesis of a period of dystrophy following the first oligo-mesotrophic period in the lake history which was disrupted and followed by a second oligo-mesotrophic period. The lake has apparently returned recently to an acidic dystrophic condition that is very similar to the earlier dystrophic period. Also of interest are the species association patterns displayed in the phenogram in Fig- ure 6. Ba.sed on the patterns of species associ- ations determined by niche overlap, it is pos- sible to infer physico-chemical requirements March 1979 Jatkar et al.: Lily Lake Diatoms 37 of some diatom taxa which are not well dis- cussed in the literature. For example, Ano- moeoneis vitrea and Nitzschia fnistulwn var. perminuta cluster at .94 niche overlap. Such a high overlap value (on a scale of 0.0- LO) indicates that their niches are highly similar. This should indicate that these two diatoms have similar ecological requirements. The same type of information can be obtained for several other species pairs as illustrated in Figure 6. pH and Saprobien Spectra The percent sum frequency of alka- liphilous, pH indifferent, and acidophilous taxa present at each level of deposition was calculated. Taxa used in this analysis are list- Nicha Ovarlap Batwaan Spacias fruftulum var. pvrminuli Nilitchio fonticula H Pinnulorio gibbo Cymb.lla ongultato Fruttulia rhomboid.. bic.p. Cymballo lunotfl rodio.. 1 Frofliiorio 30 cm tall) on more or less straight line tran.sects through C. nauseosus populations. Fifty plants were scored in each of the pure stands. At the mixed sites 50 plants of each sub- species were scored as the plants occurred on the transects. All data collection and observa- tions were made in the winter and early spring of 1976-77. Chrysothamnus nauseosus subspecies were identified using Anderson's (1973) key at the time of data collection or during the follow- ing summer when leaves were present on the plants. Herbarium vouchers have been depos- ited at the Shrub Sciences Laboratory Herba- rium (SSLP) in Provo, Utah. Flies were reared to maturity from a few galls, and were identified using Wangberg's (1976) key. A paired t test (Woolf 1968) was used to test the significance of gall form frequency differences where two gall types occurred on the same subspecies (Table 1). Results and Discussion Correlation of gall morphology with Chrysothamnus nauseosus subspecies.— The five study areas of Table 1 reflect almost complete gall form specificity for each sub- species. In each case C. nauseosus ssp. albi- caulis is growing with either C. nauseosus ssp. consimilis or ssp. graveolens. The cotton gall was found only on ssp. consimilis and ssp. graveolens. The callus gall, except for a single plant at the Paragonah .site, was found only on ssp. albicaulis. Both ssp. consimilis and s.sp. graveolens tend to have greener stems and less leaf and stem tomentum than ssp. al- March 1979 McArthur et al. : Galls on Rabbit Brush 83 Gall type Callus Cotton x±sd^ Range of values Percent of plants w/galls x±sd^ Range of values Percent of plants w/galls 7.92 ±8.55 0 0-34 0 80 0 0 0 0 0 0 0 1.00±1.76 0 0-9 0 46 0 0 0.70 ±0.90 0 0-3 0 50 4.32 ±4.83 0 0-18 0 80 0 0 4.44 ±6. 18 0 0-23 0 58 6.02 ±4.83 0.04 ±0.28 0-21 0-2 94 2 0 5.36 ±2.98 0 0-11 0 98 5.56 ±5.00 0 0-21 0 90 0 0 3.50 ±6.45 0 0-14 0 80 bicaulis. The exceptional Paragonah plant could well have been an introgressant. At the Big Rock Candy Mountain site galls were not found on the ssp. consimilis plants (Table 1). However, a few plants of intermediate mor- phology were located— they had callus galls. At the Snow Field Station the four sub- species of C. nauseosus are growing in a uni- form garden. There callus galls were found on ssp. albicaulis, and cotton galls were found, although much less frequently, on the other three subspecies. In the uniform population studies (Table 2), only callus galls were observed for C. nau- seosus ssp. albicaulis, and only cotton galls were observed for ssp. consimilis. Cotton galls were most common on ssp. graveolens, although a few mace galls were found on this subspecies at the Hanksville site. Only mace galls were observed on ssp. salicifolius. At sites where observations were made but gall frequency data were not recorded, we rarely found exceptions to the data presented in Table 2. A population of ssp. albicatilis (collection nimiber: McArthur, Tiernan, and Welch 769) located 3 km south of Nephi, Juab Co., Utah, had a single plant corre- sponding to ssp. consimilis. Whereas the ssp. albicaulis plants had high numbers of callus galls, the aberrant ssp. consimilis plant had only a single gall, which was a callus gall. Galls were much more common in some areas than in others. For example, ssp. albi- caulis plants near Ogden Bay, Weber Co., Utah (McArthur 786), were nearly devoid of galls; whereas a few miles south, on the north end of Antelope Island, Davis Co., Utah, plants of the same subspecies (McArthur 789, Table 2) were heavily infested with galls. In the upper Sevier Valley, ssp. consimilis plants lacked galls, but in the lower Sevier Valley and in Sanpete Valley (a Sevier Valley tribu- tary) they had cotton galls. A few mace galls were found with more numerous callus galls on ssp. albicaulis in Willow Creek Canyon, north of Richfield, Sevier Co., Utah. Mace galls were found on all four sub- species in at least one location of our obser- vations. The mace galls were encountered less frequently and had a lower incidence per plant than the other two types of galls. East of Ephraim, Sanpete County, Utah, we found both ssp. albicaulis (McArthur 779) and ssp. consimilis (McArthur 780) growing with several plants of intermediate morpho- logy (McArthur 781). There, callus galls were on ssp. albicaulis plants; cotton galls were on ssp. consimilis plants; and both kinds of galls were foimd on a few plants of intermediate morphology. Wangberg (1976) foimd that the tephritid fly (Aciurina bigehviae) was responsible for galls whose descriptions match those of the callus and mace forms discussed in this re- port. Another tephritid (A. maculata) was re- sponsible for galls matching the description 84 Great Basin Naturalist Vol. 39, No. 1 f-~^ ._J Fig. 1. Drawings of gall forms. A. callus. B. cotton. C. mace. (4X). March 1979 McArthur et al.: Galls on Rabbit Brush 85 of the cotton gall. Wangberg (1976) de- scribed the galls but did not use the terms cotton, callus, or mace. He (Wangberg 1976) stated that in Idaho both cotton and callus gall forms were found on C. nauseosus ssp. albicaiilis and ssp. consimilis. Observations made during this study in Utah (Tables 1 and 2) and northern Arizona were quite different. The callus gall was very specific for ssp. albi- caiilis. The cotton gall was specific for ssp. consiinilis and graveolens at most locations. Cotton galls were found on ssp. albicaiilis only where the callus gall was absent. For ex- ample, in northeastern Mohave Co., Arizona, and western Kane Co., Utah, ssp. albicaiilis, graveolens, and consimilis all had cotton galls (McArthur 819, 820, 832). Mace galls were much rarer than callus or cotton galls in most areas of observation (Tables 1 and 2), but were occasionally found on all four sub- species of C. nauseosus. The relationship between gall tomentum and plant tomentum is perplexing. Glabrous callus and mace galls have specificity for ssp. albicaiilis and ssp. salicifoliiis, respectively. These subspecies have looser and more ob- vious tomentum than ssp. consimilis and graveolens, which show specificity for the to- mentulose cotton gall. Taxonomic indicator value of gall FORMS ON C. nauseosus subspecies.— The callus gall form is specific for C. nauseosus ssp. albicaiilis in most of Utah. If a callus gall is present on C. nauseosus, the plant is in all likelihood ssp. albicaiilis (Table 1 and 2). In winter, when leaves are lacking from C. nau- seosus, callus galls help distinguish ssp. albi- caiilis from ssp. consimilis and ssp. salici- folius. In Idaho apparently, this specificity does not occur (Wangberg 1976). The cotton gall, in Utah, is found most commonly on ssp. consimilis and graveolens (Tables 1 and 2). However, in the few areas where the callus gall is missing the cotton gall may be found on ssp. albicaiilis. The cot- ton gall has not been found on naturally Table 2. Gall frequencies in uniform populations at nine Utah localities. Collection information Gall type x±sd^ Frequency Range of values Sub- species' Authors' number Site Percent of plants w/galls Chna- 768 Mouth of Chicken Creek Canyon, Juab Co. Callus 2.03 ±.3.34 0-18 72 Chna^ 773 3 km S. Bluffdale, Salt Lake Co. Callus L80±2.03 0-8 60 Chna» 789 N. end Antelope Island, Davis Co. Callus 5. 10 ±7.26 0-43 74 Chna^ 962 Pigeon Hollow, Sanpete Co. Cotton L24±1.41 0-6 68 Chna'^ 976 3 km NE. Ephraim, Sanpete Co. Cotton 1.06 ±1.05 0-4 68 Chnag 1000 Hanksville, Wayne Co.' Cotton' 1.44 ± 1.69 0-7 66 Chna^ 964 Snowberry Exclosure, Ephraim Canyon, Sanpete Co. Mace 0.18 ±0.52 0-3 • 14 Chna^ 1061 Red Creek, Duchesne Co. Mace^ 0.26 ±0.49 0-2 24 Chna^ 1066 Moon Lake, Duchesne Co. Mace 0.16 ±0.55 0-3 10 'Subspecies symbols: Chna^ = Chrysothamnus nauseosus ssp. albicaulis; Chna* = C. n. ssp. consimilis. ChnaS = C. n. ssp. graveolens; ssp. salicifolius. 'x± = mean (x) ± standard deviation (sd) of number of galls on terminal 15 cm of a random branch/plant. Two morphologies of ssp. graveolens present. A low incidence of mace galls was observed but not recorded. 'One poorly defined cotton gall was observed. ChnaS = C. n. Great Basin Naturalist Vol. 39, No. 1 growing populations of ssp. salicifolius, but was found on plants of this subspecies trans- planted to the Snow Field Station. Subspecies salicifolius is ordinarily a mountain taxon, whereas the Snow Field Station is in a valley setting. The mace gall has been found on all four subspecies but is the only one of the three galls found on naturally occurring popu- lations of ssp. salicifolius (Table 2). Resource partitioning by the gall- forming TEPHRITID FLIES.— Waugbcrg (1976) discussed the problem of niche-sharing by fly species that induce galls on similar parts of the same subspecies of C. nauseosus at the same time of the year. Wangberg be- lieved, following Darlington (1972), that tephritid fly species compete in nature and divide the resource. Our studies support such a competitive division of the plant resource. Our study showed more gall-form specificity on C. nauseosus subspecies than Wangberg's (1976), perhaps because our study was in the central area of C. nauseosus's range, where there are larger concentrations of the plant than are present where Wangberg studied (Hall and Clements 1923). L. C. Anderson (letter dated 16 February 1978) has data in- dicating less specificity for gall forms on plants growing outside of Utah than on those growing in Utah. In the central area of the host species' occurrence, the gall formers might be better off to be host specific to avoid competition with large populations of various taxa of gall-forming flies. In outlying areas such host specificity might be a dis- advantage because host plants would be rarer. Moreover, gall-forming fly populations would not be as high, so competition would be reduced. MacArthur (1972:17) touched on this problem when he stated that species are more likely to compete in localities where an advantage can be gained. Another possible explanation for gall-form specificity is that different gall forms are not a re.^ponse to different fly species but rather a response to the host plant species. We have not reared enough fly specimens to address this possibility. However, we think this expla- nation is unlikely, because a few individual plants had more than one gall form. Acknowledgments This study was facilitated by federal funds for wildlife habitat restoration made avail- able from the Pittman-Robertson W-82-R Project (Cooperators: Intermountain Forest and Range Experiment Station, USDA Forest Service, Ogden, Utah; and Utah State Divi- sion of Wildlife Resources, Salt Lake City, Utah). The Snow Field Station is coopera- tively maintained by the above agencies, Utah State University, and Snow College. The drawings of Figure 1 were made by An- nielane Jones Yazzie. Drs. L. C. Anderson, D. L. Hanks, K. T. Harper, and J. K. Wangberg provided technical advice. Literature Cited Anderson, L. C. 1966. Cytotaxonomic studies in Chrysothamnus (Astereae, Compositae). Am. J. Bot. 53:204-212. 1971. Additional chromosome numbers in Chrysothamnus (Asteraceae). Bull. Torrev Bot. Club 98:222-225. 1973. Chrysothamnus, p. 109-111. In: S. L. Welsh and G. Moore. Utah Plants. 3d ed. Brig- ham Young Univ. Press, Provo, Utah. 474 p. Anderson, L. C, D. W. Kyhos, T. Mosquin, .\. M. Powell, and P. H. Raven. 1974. Chromosome numbers in Compositae. IX. Happlopappus and other Astereae. Am. J. Bot. 61:665-671. Branson, F. A., R. F. Miller, and I. S. McQueen. 1976. Moisture relationships in twelve northern desert shnib communities near Grand Junction, Colo- rado. Ecology 57:1104-1124. Cronquist, a. 1955. Part 5: Compositae. In: C. L. Hitchcock, A. Cronquist, M. Owenby, and J. W. Thompson. Vascular plants of the Pacific North- west. Univ. Washington Press, Seattle, Washing- ton. 343 p. Cronquist, A., A. H. Holmgren, N. H. Holmgren, and J. L. Reveal. 1972. Intermountain flora, vascular plants of the Intermountain West. U.S..\. Vol. I. Hafner Publ. Co., New York, New York. 270 p. Darlington, P. J., Jr. 1972. Competition, competitive repulsion, and coexistence (fainial equilib- riums/exclusion principle/niche/ecologic range). Proc. Natl. Acad. Sci. USA 69:3151-3155. Hall, H. M., and F. C. Clements. 1923. The phyloge- netic method of taxonomy. The North American species of Artenmia, Chrysothamnus, and Atri- plex. Carnegie Inst. Publ. 326:1-355. Hanks, D. L., E. D. Mc.\rthur, \. P. Plummer, B. C. GiuNTA, and a. C. Blauer. 1975. Chromatogra- phic recognition of some palatable and impala- table subspecies of rubber rabbitbrush in and around Utah. J. Range Manage. 28:144-148. March 1979 McArthur et al.: Galls on Rabbit Brush 87 MacAhthur, R. H. 1972. Geographical ecology: patterns in the distribution of species. Harper and Row Publishers, New York, New York. 269 p. McArthur, E. D., A. C. Blauer, A. P. Plummer, and R. Stevens. 1979. Characteristics and hybridiza- tion of important intermountain shrubs. III. Sun- flower family. USDA For. Serv. Res. Pap. INT 220. Intermt. For. and Range Exp. Stn., Ogden, Utah. 82 p. McArthur, E. D., B. C. Giunta, and A. P. Plummer. 1974. Shnibs for restoration of depleted ranges and disturbed areas. Utah Sci. 35:28-33. McArthur, E. D., D. L. Hanks, A. P. Plummer, and A. C. Blauer. 1978. Contributions to the taxonomy of Chrysothammis viscidiflorus (Astereae, Com- positae) and other Chrysothammis species using paper chromatography. J. Range Manage. 31:216-223. Plummer, A. P. 1977. Revegetation of disturbed inter- mountain area sites, p. 302-339. In: J. L. Thames (ed.) Reclamation and use of disturbed land in the Southwest. Univ. Arizona Press, Tucson. 362 P- Wangberg, J. K. 1976. Biology of the tephritid gall- formers. (Diptera: Tephritidae) on rabbitbrush, Chrysothatnnus spp., in Idaho. Unpublished Ph.D. Dissertation, Univ. Idaho, Moscow. 240 p. (University Microfilms, Ann Arbor, Michigan. Order No. 77-6706). WooLF, C. M. 1968. Principles of biometry. D. Van Nos- trand Co., Inc. Princeton, New Jersey. 359 p. HOMING BY A PYGMY RABBIT' Jeffrey S. Green^'^ and Jerran T. Flinders' Abstract.— A juvenile female pygmy rabbit {SylviUigtis idahoensis) escaped from a holding pen and was recap- tured 211 days later 200 m from its original capture site and 2.5 km from the pen facility. Homing for distances of 3.8 km (Hill 1967) and 4.8 km (Bowers 1954) was reported for the eastern cottontail {Sylvilagus floridanus). Chapman (1971) reported the brush rabbit (S. bachmani) failed to home greater than 0.16 km in 29 trials. This is a report of homing by a pygmy rabbit (S. idahoensis), the smallest member of the genus, from a distance of 2.5 km. The rabbit, a juvenile female weighing 101 g, was captured 23 June 1977 on the U.S. Sheep Experiment Station near Dubois, Idaho. It was taken to a permanent pen facil- ity, ear-tagged, and placed in a cage from which it escaped during the first night. Two hundred and eleven days later on 20 January 1978, while trapping pygmy rabbits for be- havioral research, the escaped specimen was caught approximately 200 m from its original capture site and 2.5 km from the pen facility. It appeared to be in good condition and it weighed 392 g. The rabbit was observed at least four times during the next 19 days in the second capture area and was apparently a resident there. Suitable habitats occupied by pygmy rabbits are found in any direction of escape from the cage area. The fact that the animal traveled toward its original capture site is, therefore, not apparently related to the absence of suitable habitat elsewhere. Maximum movements we previously re- corded for two pygmy rabbits (tracked in snow) were one-way distances of 140 and 450 m during winter 1976. Wilde et al. (1976) noted an adult female (radio-instrumented in late November) that showed extreme fidelity to her burrow and was never located farther than 20 m from the burrow where originally captured. However, they documented longer movements by tagged and recaptured indi- viduals; one female moved a maximum of 300 m, a male moved as far as 500 m, and an- other male moved 200 m. Janson (1946) ob- served that activity of pygmy rabbits in win- ter was within a radius of about 27 m from their burrows. He wrote that in spring the radius of activity increased in response to snow melt and onset of breeding activity. On the basis of McNab's (1963) method of pre- dicting approximate size of home range from basal metabolism and body size of mammals, the home range of the pygmy rabbit should be about 0.8 ha. This was calculated using an average body weight of 454 g obtained from 11 female pygmy rabbits kept in captivity and weighed at least weekly from September through February. Clearly the movement we recorded is far beyond that normally associ- ated within a radius of movement well within documented range for the species. A move- ment of 2.5 km for a juvenile sheds light on the dispersal and pioneering capabilities of the pygmy rabbit. Literature Cited Chapman, J. A. 1971. Orientation and homing of the bnish rabbit. {Sylvilagus bachmani). J. Mamm. 52:686-699. Bowers, G. L. 1954. An evaluation of cottontail rabbit management in Pennsylvania. Trans. N. Amer. Wildl. Conf. 19:358-367. Hill, E. P. 1967. Homing by a cottontail rabbit. J. Mamm. 48:648. Janson, R. G. 1946. A survey of the native rabbits of Utah with reference to their classification, distri- bution life histories and ecology. Unpublished master's thesis. Utah State University, 103 pp. McNab, B. K. 1963. Bioenergetics and the determina- tion of home range size. Amer. Nat. 97:133-140. Wilde, D., J. S. Fisher, and B. L. Keller. 1976. A de- mographic analysis of the pygmv rabbit, Sy/- vilagus idahoensis. Pp. 88-105, In: 1975 Progress Report Idaho National Engineering Laboratory Site Radioecoiogy-Ecology Programs (O. D. Mar- ham, ed.), U.S. Energy Res. and Development Admin., Idaho Falls. 205 pp. 'This study was supported by USDA Western Region Federal Cooperative Agreement 12-14-5001-264. 'Department of Botany and Range Science, Brigham Young University, Prove, Utah 84602. 'Present address: U.S. Sheep Experiment Station, D\ibois, Idaho 8.342,3. 88 BEETLES FROM THE ENVIRONS OF LAKE POWELL IN SOUTHERN UTAH AND NORTHERN ARIZONA Dorald M. Allred' and Vasco M. Tanner' Abstract.— Sixty-eight species of beetles of 14 families were collected in pit traps in 12 major vegetative commu- nities during environmental monitoring studies in the summers of 1971 to 1973 in southern Utah and northern Ari- zona. Seasonal and plant community differences and correlations in population and composition were noted for the 16 sites studied. Highest populations and the most species were foimd in a grass commimity, but occurred in differ- ent months in each of the three years. Lowest populations were found in a Coleogyne community, and fewest species in an A rtemisi« -grass community. In June 1971 an ecological study was in- itiated by the Center for Health and Environ- mental Studies at Brigham Young University to establish baselines to determine the envi- ronmental effects of the Navajo Generating Station near Page in northern Arizona, and the proposed Kaiparowits Generating Station in southern Utah.- Field studies of arthropods were conducted from July to September in 1971, and from May to August in 1972 and 1973. Can pit traps (an outer galvanized metal sleeve 18 cm in diameter and 36 cm long with a stainless steel, flanged inner can of slightly smaller size) were used to capture ground-dwelling arthropods. Five traps 30 m apart were placed on each of two transects, which were 45 m apart. These were left open, dry, and unbaited for a 72-hour period once each summer month. Of the 12 sites studied in 1971, the 16 in 1972, and the 12 in 1973, some were studied only one year, oth- ers two, and some all three years. Beetles, other arthropods, small rodents, and lizards were collected from the cans daily while they were open for trapping studies. The beetles were identified by Dr. Vasco M. Tanner. Study Sites and Their Predominant Vegetation Site 1. Ephedra-Vanclevea-Sporobolus-Ory- zopsis-Hilaria. Base N slope Cedar Mtn, 5 km W Glen Cnyn City, Kane Co., Utah. Site 2. Juniperus-Ephedra-Muhlenbergia- Botiteloua-Hilaria-Oryzopsis. Cedar Mtn, 6.5 km S site 1. Site 3. Ephedra-Hilaria-Bouteloua-Ory- zopsis. Cedar Mtn, 2 km S site 2. Site 4. Coleogyne-Ephedra-Atriplex-Chryso- thamnus. Cedar Mtn, 1.3 km SE site 3. Site 6. Artemisia-Hilaria-Aristida-Ory- zopsis. Smokey Mtn, 23 km from Last Chance Jnct, Kane Co., Utah. Site 8. Grayia-Ephedra-Coleogyne-Hilaria- Bouteloua-Oryzopsis. Smokey Mtn, 14.5 km from Last Chance Jnct., Kane Co., Utah. Site 10. Ephedra-Yiicca-Eurotia-Vanclevea- Oryzopsis-Streptanthella. Ahlstrom Pnt Rd., 7.5 km S junct., Kane Co., Utah. Site 13. Grayia-Ephedra-Oryzopsis-Bou- teloua-Hilaria. Nipple Bench, 6.5 km SE Tib- bet Spring, Kane Co., Utah. Site 14. Coleogyne-Grayia-Ephedra- Chrysothamnits-Hilaria. 3 km S site 13. Site 19. Coleogyne. 15 km S Page, Coco- nino Co., Arizona by Hwy 89. Site 20. Muhlenbergia-Bouteloua-Hilaria. 19 km S Page, Coconino Co., Arizona by Hwy 89. Site 22. Coleogyne-Ephedra-Hilaria. E Navajo Generating Station, SE Page, Coco- nino Co., Arizona. Site 23. Ephedra-Coleogyne-Grayia-Hi- laria. Cathys Flat, 2 km N Tibbet Spring, thence 2.5 km E, Kane Co., Utah. Site 27. Juniperiis-Pinus. Four-mile Bench, 5 km SE cow camp, head Wesses Cnyn, Kane Co., Utah. 'Department of Zoology and Life Science Museum. Brigham Young University, Provo, Utah 84602. 'These studies were funded cooperatively by Arizona Pubhc Service, Los Angeles Power and Light, Salt River Project, San Diego Power and Light, and Southern California Edison. 89 90 Great Basin Naturalist Vol. 39, No. 1 Site 28. Artemisia-Bouteloua-Plantago. 3 km E site 27. Site 30. Ephedra- Bouteloua-H ilaria-Sporo- holus-Salsola. Brigham Plains Flat, Kane Co., Utah. Annotated List of Families and Species Buprestidae Acamaeodera lanata Horn: one specimen, 17 July 1972, site 14 {Coleogyne-Grayia-Eph- edra-graiss); 6 specimens, 2 June 1973, site 20 (grass). Carabidae Bembidion sp.: one specimen, 13 May 1973, site 27 (juniper-pinyon). Calosoma triste Lee: one specimen, 3 May 1973, site 8 (Grat/ia-grass). Calosoma sp.: one specimen, 5 July 1973, site 10 (Ephedra-grass). Celia sp.: one specimen, 5 June 1972, site 3 (Ephedra-grass); one specimen 6 June 1972, site 1 (Ephedra-Vanclevea-grass). Harpahis corpulentus (Csy.): 6 specimens were taken 4 June 1973 at site 3 [Ephedra- grass); 2 specimens 6 June 1973, at site 8 (Grai/ia -grass); and one specimen 9 July 1973 at site 14 (Coleogyne-Grayia-Ephedra-grass). The apparent absence of this species in 1972 is unusual, inasmuch as sites 3, 8, and 14 were studied from May through August. Its ab- sence in 1971 is also unexpected, even though collections were made only in July and Au- gust. Harpahis fitrtivus (Lee): 3 specimens, 21 August 1972, site 28 (Artemisia-grass); one specimen, 5 September 1972, site 3 (Ephedra- grass). Harpahis histraiis Csy.: 2 specimens, 4 June 1973, site 3 (Ephedra-grass); one speci- men, 14 June 1973, site 28 (Arterjiisia-grass). Harpahis sp.: one specimen, 10 July 1973, site 23 (Ephedra-Coleogyne-Grayia). Rhadine jejuna Lee: one specimen, 12 June 1972, site 19 (Coleogyne). Rhadine sp.: one specimen, 4 July 1973, site 8 (Grai/ia -grass). Selenophorus aeneopiceus Csy.: 3 speci- mens, 12-13 June 1972, site 19 (Coleogyne); one specimen, 22 June 1972, site 30 (Eph- edra-grass). Cerambycidae Moneilema obtusa Lee: one specimen, 1 August 1973, site 10 (Ephedra-grass). Prionus sp.: one specimen, 7 July 1973, site 30 (Ephedra-grass). Chrysomelidae Disonycha fiimata (Lee): 4 specimens, 12- 13 July 1971, site 3 (Ephedra-grass). Disonycha latifrons Schaffr.: one speci- men, 3 May 1973, site 6 (Artemisia); one specimen, 6 May 1973, site 30 (Ephedra- grass). Cleridae Cymatodera fuchsi Schaffr.: one specimen, 12 August 1971, site 13 (Grayia-Ephedra- grass); one specimen, 10 July 1972, site 10 (Ephedra-grass). Coccinellidae Hippodamia convergens Guer.: one speci- men, 2 June 1973, site 20 (grass). Colydiidae Bitoma sp.: one specimen, 26 September 1971, Glen Cnyn City. Curculionidae Cimbocera huchanani Ting: 3 specimens, 13-14 July 1971; one specimen, 10 November 1971, site 3 (Ephedra-grass); one specimen, 15 October 1971; one specimen, 9 May 1973, site 14 (Coleogyne-Grayia-Ephedra-grass); 3 specimens 14-15 July 1972; one specimen, 14 August 1972, site 20 (grass); one specimen, 10 May 1973, site 23 (Ephedra-Coleogyne-Gray- ia). Cimbocera conspersa Fall: 4 specimens were taken at site 2, one at site 3, one at site 6, 4 at site 14, 2 at site 23, and 12 at site 27. Plants of the genus Ephedra were common to four of the six sites where this beetle was found. However, greatest numbers were found in the juniper-pinyon community. Five specimens were taken in April and May 1972, and 19 from May to August 1973. Greatest numbers were found in May of both years. The activity of this .species apparently is spring and early .summer, for no specimens were taken in July or August of 1971, none after May in 1972, and only four after May in 1973. March 1979 Allred, Tanner: Ecology of Beetles 91 Cimbocera petersoni Tanner: one speci- men, 29 April 1973, site 2 (juniper-Ephedra- grass); one specimen, 30 April 1973, site 1 {Ephedra- Va nc/eoea -grass) . Dinocleus angularis (Lee): one specimen, 4 May 1973, site 10 (Ephedra-grsLSs). Eucyllus unicolor Van Dyke: one speci- men, 5 August 1972, site 3 (Ephedra-grsiss). Eupagoderes varius (Lee): one specimen, 12 July 1972, site 6 (Artemisia). Eupagoderes sp.: one specimen, 3 June 1973, site 2 (juniper-£p/iedra -grass). Ophryastes sordidus Lee: one specimen, 9 September 1971, site 7 (Grayia-Coleogyne- grass). Ophryastes sulcirostris (Say): one specimen, 4 June 1973, site 2 (juniper-Ep^edra-grass). Ophryastes sp.: one specimen, 13 May 1973, site 28 (Artemisia-grsiss). Histeridae Saprinus discordalis (Lee): one specimen, 20 July 1971, Navajo Mtn. Saprinus lugens Et.: 6 specimens were taken at site 2, 4 at site 10, one at site 14, and 23 at site 22. Plants of the genus Ephedra were common to three of the four sites where this species was found. However, greatest numbers were found in the Coleogyne com- munity. Twenty-three specimens were taken in July 1972, and 5 and 6 in May and July of 1973, respectively. Apparently July is the month of greatest activity of this species, even though none were taken in July of 1971. However, site 2 was the only one of the four sites where this species was found that was studied during July of 1971. Saprinus oregonensis Lee: 2 specimens, 2 August 1971, site 6 (Artemisia). Meloidae Pyrota mylabrina Cher.: 34 specimens, 24 August 1972, Cottonwood Cnyn, 9 mi N Hwy 89, ex Chrysothamnus. Melyridae Collops femoratus (Schaffr.): one specimen, 14 July 1971, site 15 (Popuhis-Tamarix-Ly- _cium). Trichochrons zionicus Tanner: 28 speci- mens, summer 1972, specific locality un- known. Ptinidae Ptinus sp.: 8 specimens were taken at site 1, one at site 2, 5 at site 14, 2 at .site 17, 6 at site 22, and 8 at site 23. Plants of the genus Ephedra and grasses were common to four of the six sites. Significant differences between community types were not evident, although greatest numbers were found in those where Coleogyne was a predominant. Four speci- mens were taken in July and August 1971, 18 from April to August 1972, and eight in May and July of 1973. Greatest numbers were found in May of 1972 and 1973. Salpingidae Conotus lanchesteri Van Dyke: This was one of the more common beetles. Twenty- one were taken at site 1, 38 at site 2, one at site 7, 6 at site 9, one at site 13, 2 at site 19, 6 at site 22, 3 at site 23, and 2 at site 27. The plant Ephedra was common to four of the nine sites, and greatest numbers were found on sites 1 and 2, where it was present. Speci- mens were taken in July and August 1971, April to September in 1972, and May to Au- gust 1973. In 1971 about equal numbers were taken in July and August, in 1972 greatest numbers were found in August, and in 1973 greatest numbers in July and August. No sig- nificant differences were noted between the three years, although slightly more numbers were taken in July and August of 1972 than during those months in other years, and more were taken from May to August in 1972 than for comparative months of 1973. Scarab aeidae Aphodius rubidus (Lee): 4 specimens were taken at site 2, 13 at site 3, 3 at site 6, 6 at site 8, 6 at site 10, 4 at site 13, 31 at site 14, 2 at site 20, 3 at site 22, and 17 at site 30. The plant Ephedra was common to 6 and grasses to 8 of the 10 sites where this beetle was found. Greatest numbers were found on sites 14 and 30, where Ephedra and grasses were common. Specimens were taken in April and from June to September in 1972, and from May to July in 1973. Greatest numbers were found in April of 1972 and May of 1973. From May to August, populations were 24 times higher in 1973 than in 1972. Diphtaxis subangulata (Schaffr.): 2 speci- mens, 16 June 1972, site 13 (Grayia-Ephedra- 92 Great Basin Naturalist Vol. 39, No. 1 grass); one specimen, 21 July 1972, site 27 (juniper-pinyon). Diplotaxis sp.: one specimen, 6 April 1971; one specimen, 6 June 1971, site 3 {Ephedra- grass); one specimen, 5 June 1971, site 1 (Ephedra-Vanclevea-grass). Polyphylla diffracta (Csy.): 3 specimens, 20 July 1971, Navajo Mtn. Polyphylla 10-lineata (Say): one specimen, 12 June 1973, site 23 {Ephedra-Coleogyne- Grayia). Serica sp.: one specimen, 11 June 1973, site 13 (Grayia-Ephedra-grsiSs); 4 specimens, 19 June 1972; 5 specimens, 20 June 1972, site 27 (juniper-pinyon). Tenebrionidae Araeoschizus dicipiens (Horn): one speci- men, 7 April 1972, site 1 {Ephedra-Van- clevea-grass); one specimen, 13 June 1972, site 19 (Coleogyne); one specimen, 9 August 1973, site 30 (Ephedra-grsiss). Chilometopon abnorme Horn: 13 speci- mens were taken at site 1, one at site 2, 9 at site 4, one at site 6, 2 at site 7, one at site 8, 56 at site 10, 2 at site 13, 4 at site 14, 8 at site 17, and 2 at site 18. These beetles were taken from a variety of vegetative types, but signif- icantly greater numbers were found at site 10, an Ephedra-grsiss community. Specimens were taken from July to September in 1971, May to August in 1972, but only in August of 1973. Beetles in July and August of 1971 were twice as abundant as in 1972, and 48 times as abundant as in 1973. From May to August, they were 30 times as abundant in 1972 as in 1973. Coniontis opacus Horn: one specimen, 13 July 1971, site 4 (Coleogyne). Coniontis sp.: one specimen, 6 May 1972, site 4 (Coleogyne). Edrotes leechi Doyen: one specimen, 4 June 1972, site 2 (juniper-Ep^edra-grass); one specimen, 14 July 1971, site 3 (Ephedra- grass); one specimen, 12 July 1971, site 4 (Co- leogyne); one specimen, 11 July 1972, site 6 (Artemisia); one specimen, 19 July 1971, site 18 (Coleogyne); 2 specimens, 14 August 1972, site 20 (grass); 2 specimens, 12 April 1973; one specimen, 12 May 1972, site 22 (Coleo- gyne). Eleodes caiidifera Lee: 7 specimens were taken at site 1, 2 at site 3, 6 at site 10, 7 at site 13, 3 at site 14, and one at site 19. Eph- edra and grasses were predominant plants at 5 of the 6 sites. No site had significantly greater numbers of beetles than another. Specimens were taken in July and August 1971, May to July 1972, and May and June 1973. In 1973, greatest numbers were found in May, but no significant differences were noted between months of other years except from May to August, when eight times as many beetles were found in 1973 as in 1972. Eleodes extricata (Say); Beetles of this spe- cies were some of the most common and widespread of those taken. A total of 164 specimens was taken from 15 of the 19 sites studied. Greatest numbers were found at sites 3, 19, 20, and 22, which are predominantly Coleogyne or grass habitats. Specimens were taken in July and August of 1971, April to September in 1972, and May to August in 1973. Greatest numbers were taken in August and September of 1972, and July of 1973. In July and August of 1972 and 1973 these bee- tles were about equally abundant, but only one-tenth as abundant in 1971. For the peri- od of May to July, they were about equally as abundant in 1972 as in 1973. Eleodes hispilahris sculptilis Blais.: Beetles of this species were common and widespread; 105 were taken from 13 study sites. Greatest numbers were found at site 10, an Ephedra- grass community. In 1971 they were found from July to September, most abundantly during the latter month; in 1972 during April and from June to September, also most abun- dantly during September; and in 1973 from May to August, mostly in July. In July and August, they were half again as abundant in 1971 as in 1972 and 1973. They occurred in about equal numbers for the period of May to August in 1972 and 1973. Eleodes immunis interstitralis Blais.: one specimen, 18 July 1971, site 17 (grass); one specimen, 19 July 1971, site 1 (Ephedra- Vanclevea-grsLSs) . Eleodes leechi Tanner: one specimen, 20 July 1972; one specimen, 22 July 1972, site 28 (Artemisia-gruss); one specimen, 11 July 1973, site 27 (juniper-pinyon); one specimen, 3 August 1973, site 14 (Coleogyne-Grayia- Ephedra -grass); 2 specimens, 4 August 1973, site 2 (juniper-Ephedra-grass). Eleodes obscura sulcipennis Mann.: These March 1979 Allred, Tanner: Ecology of Beetles 93 beetles were relatively abundant and wide- spread and were found on 15 sites. They were most abundant at sites 2, 3, 8, 14, and 19, principally where grasses were pre- dominant. Beetles were taken from July to September, mostly August in 1971; from April to September in about equal numbers each month in 1972; and from June to Au- gust, also in about equal numbers each month in 1973. In July and August, populations were three times as high in 1971 as in 1972 and 1973. From May to August about equal numbers were taken in 1972 and 1973. Eleodes omissa pygmaea Blais.: 14 speci- mens, July and August 1971, site 3 {Ephedra- grass); one specimen, 1 August 1971, site 6 (Artemisia); one specimen, 11 August 1971, site 13 (Grayia-Ephedra-gTSLSs); 2 specimens, 11, 13 August 1971, site 14 (Coleogyne-Gray- ia-Ephedra-gr ass); one specimen, 18 July 1971; 2 specimens, 27 August 1971, site 17 (grass); one specimen, 17 July 1971; one spec- imen, 17 August 1971, site 18 (Coleogyne). Eleodes porcata Csy.: one specimen, 8 Au- gust 1973, site 39 {Ephedra-gYa.ss). Efnbaphion glabrum Blais.: 3 specimens were taken at site 1, 3 at site 2, 6 at site 3, 2 at site 19, 4 at site 22, one at site 23, 17 at site 27, and 14 at site 28. Greatest numbers were found at sites 27 and 28. Beetles were taken in July and August of 1971, April to September of 1972, and May, July, and Au- gust of 1973. During July and August about equal numbers were found in each of the three years. For the period of May to August, twice as many beetles were taken in 1972 as in 1973. Exchatomoxys tanneri Sorenson and Stones: one specimen, 15 August 1971, site 3 (Ephedra-graiss). Glyptasida sordida (Lee): one specimen, 13 August 1971; one specimen, 17 August 1972, site 13 (Grayia-Ephedra-grass); one specimen, 12 August 1971; one specimen, 16 July 1972, site 14 (Coleogyne-Grayia-Eph- edra-grass); 3 specimens, 13-14 August 1972, site 19 (Coleogyne); 24 specimens, July and August 1972, site 20 (grass). Glyptasida sp.: one specimen, 15 August 1971, site 3 (Ephedra-grass). Hylocriniis delicatulus Csy.: 6 specimens were taken at site 1, 2 at site 14, 9 at site 19, 2 at site 20, and 18 at site 22. Most specimens were found at sites 19 and 22, which are Co- leogyne communities. Beetles were taken from June to August in 1972, and May to Au- gust in 1973. None were found in 1971. In 1972, highest numbers were taken in July, but populations were not significantly differ- ent between months in 1973. About twice as many beetles were taken in 1972 as in 1973. Metaponium sp.: one specimen, 17 July 1972, site 14 (Coleogyne-Grayia-Ephedra- grass). Pelecyphoms haruspex Csy.: 2 specimens, 28 August 1971, site 1 (Ephedra-Vanclevea- grass); 2 specimens, 6 August 1972; one spec- imen, 6 September 1971, site 2 (juniper-Ep/i- edra-grass); 4 specimens, 14 August 1971; one specimen, 25 August 1971; one specimen, 1 August 1973, site 3 (Ephedra-grass); 5 speci- mens, 18, 20 August 1971; 7 specimens, 11-12 August 1972, site 10 (Ephedra-grass); one specimen, 28 August 1971, site 18 (Coleog- yne). Sphaeriontis muricata Lee: 65 beetles were taken from 11 study sites. They were most abundant on sites 10 (Ephedra-grass), 19, and 22 (Coleogyne). Specimens were taken in July and August of 1971, May to September of 1972, and in May, June, and August of 1973. Greatest numbers in 1972 were found in July, but in 1971 and 1973 the populations were about equal each month. In July and August twice as many beetles were taken in 1972 as in 1971, and five times as many as in 1973. From May to August four times as many beetles were taken in 1972 as in 1973. Steriphanus sp.: one specimen, 7 April 1972, site 3 (Ephedra-grass). Triorophus lariversi Blais.: one specimen, 14 August 1971, site 2 (juniper -Ephedra- grass). Triorophus tenehratulus Csy.: 77 beetles were taken from 10 sites, most abundantly from site 22, a Coleogyne community. Only two specimens were found in August 1971; most were taken from April to August in 1972, and May to August in 1973. In 1972 and 1973, greatest numbers were found in May. From May to August, only a few more specimens were taken in 1972 than in 1973. Trogloderus tuberculatus Blais: Beetles of this species were the most abundant and widespread of all species collected. A total of 94 Great Basin Naturalist Vol. 39, No. 1 510 specimens was taken from all 19 study sites. Greatest numbers were found on site 19, a Coleogyne community. Specimens were found every month that field studies were conducted. In 1971, greatest numbers were found in July, and in 1972 and 1973 in June. In July and August, from two and one-half to three times as many beetles were taken in 1972 as in either 1971 or 1973. From May to August, half again as many beetles were taken in 1972 as in 1973. Discussion An initial objective was to select study areas in the major vegetative types within a 48-km perimeter of the proposed sites of the electric generating stations. Retention of study plots beyond the first year was based on vegetative type, direction from the poten- tial source of pollution, and especially the species and relative abundance of organisms present that could be used as indicator spe- cies to monitor environmental changes. Some sites were discontinued after one season be- cause of inaccessibility, discontinuous vegeta- tive analyses, and specific climatic and eda- phic data. In order to compare populations and sea- sonal changes, the numbers of beetles collect- ed were adjusted to the number of trapping attempts. The normal variability in seasonal and annual populations, slightly different dates of collection each month, and the in- fluence of periodic and abrupt climatic changes on the activity of the beetles during the trapping periods were ignored. Never- theless, the method used adequately deter- mined relative abundance and distribution within the limitations of time, economy, and logistics. The number of trapping attempts is shown in Table 1 . Pit traps are effective primarily for ground-dwelling arthropods that move on the ground more frequently than they fly. The traps involve minimum effort and time and can be used effectively for those species which may be so trapped. In this study about half of the beetles caught were in the families Carabidae and Tenebrionidae, whose mem- bers are primarily ground-dwellers. Nineteen of the species collected were sufficiently abundant to be used as indicators, and 14 of these belonged to the Tenebrionidae, the darkling beetles (Table 2). Highest populations of beetles of all spe- cies occurred in August 1971, July 1972, and June 1973 (Table 3). These annual differences likely are related to climatic and vegetational variations, and when such data are available, multiple regression analyses may show specif- ic correlations. Table 1. Number of trap-days' for pit traps on 16 major study sites, 1971-1973. Site 1971 1972 1973 Total 1 60 120 90 270 2 60 180 90 330 3 60 180 90 330 4 60 120 " 180 6 60 90 90 240 8 60 90 90 240 10 30 120 90 240 13 60 150 90 300 14 60 150 90 300 19 90 ° 90 20 90 ° 90 22 150 ' 150 23 120 90 210 27 120 70 190 28 120 70 190 30 150 90 240 Total 510 2,040 1,040 3,590 'Niimber of traps multiplied by number of days operated. 'Not operated during year indicated. Table 2. Numbers and distribution of beetles' on 16 major study sites, 1971-1973. Total No. sites number where Species taken found Trogloderus tuherculatus 510 16 Eleodes sulcipennis 192 12 Eleodes extricata 164 14 Eleodes hispilabris sculptilis 105 10 Chilometopon abnomie 100 8 Aphoditis rubidus 89 10 Conotus lanchesteri 80 7 Eleodes obscura sulcipennis 78 7 Triorophus tenebratulus 77 9 Sphaeriontis muricata 65 11 Embaphion glahrum 50 6 Htjlocrinus delicatulus 37 6 Sapriniis Uigens 34 4 Glyptasida sordida 31 4 Ptinus sp. 30 5 Eleodes caudifera 26 6 Pclect/phorus haruspex 25 4 Cimbocera conspersa 24 6 Eleodes omissa pygmaea 23 4 'Only those species are included of which more than : taken. March 1979 Allred, Tanner: Ecology of Beetles 95 The number of species of beetles which oc- curred on each study site was not consistent for each of the three years (Table 4). Even between sites with the same predominant species of plants, variations occurred be- tween sites and years. This is expected be- cause the presence, abundance, and vigor of plants, especially annual grasses and forbs, varies from year to year as influenced by the local climate. Comparison of these sites showed some significant differences. Sites 4, 19, and 22, primarily Coleogyne, had only one species of darkling beetle in common. Sites 19 and 22 had three additional species in common. Site 4 had two species not found on the other two sites, site 19 had two unique species, and site 22 had one unique species. Sites 14 and 23, which also contain Coleo- gyne as a predominant shrub, had one species in common with the other Coleogyne sites. £p/ief/r«-grassland sites 3, 10, and 30 had no species in common. One species was com- mon to sites 10 and 30, one to sites 3 and 30, and one to sites 3 and 10. Three species were unique to site 3, and four to site 10. Com- parison of sites 1 and 20 (which also had grass as a predominant component) with sites 3, 10, and 30 showed one species in common, one common to sites 1, 3, and 30, one com- mon to sites 1 and 10, one common to sites 3 and 20, and one species unique to site 20. Artemisia sites 6 and 28 had one species in common, two unique for site 6, and one unique to site 28. Grayia-grsLSS sites 8 and 13 had two species in common, and two unique to site 8. Juniper woodland sites 2 and 27 had one species in common, two unique to site 2, and one unique to site 27. Species composition differed during the Table 3. Total number of beetles of all species collected in pit traps at 16 major study sites, 1971-1973. No. beetles collected Month 1971 1972 1973 Actual Adjusted' Actual Adjusted' Actual Adjusted' April o 88 141 ' May • 74 111 192 250 June • 208 250 121 290 July 83 332 367 367 118 236 August 232 394 239 287 114 228 September 25 200 49 157 ° 'Numbers adjusted to number of trap days. "Traps not operated. Table 4. Number of species captured in pit traps on 16 major study sites, 1971-1973. Site Predominant vegetati 1971 1972 1973 Total 1 Ephedra-Vanclevea-Grass 2 Junipenis-Ephedra-GTSiSS 3 £p/iedra-Grass 4 Coleogyne 6 Arternisia 8 Grayia -Grass 10 Ephedra-Grass 13 Grayia-Ephedra-Grass 14 Coleogyne-Grayia-Ephedra-Grass 19 Coleogyne 20 Grass 22 Coleogyne 23 Ephedra-Coleogyne-Gratjia 27 Juniper-Pinyon 28 Artemisia-Grass 30 Ephedra-Gxass 9 12 11 18 6 10 12 20 14 11 11 26 7 6 • 9 5 8 5 13 3 4 8 9 6 9 10 15 9 8 15 13 15 20 13 • 13 9 • 9 11 • 11 8 9 12 5 7 10 4 5 7 4 7 10 *Not trapped during year indicated. 96 Great Basin Naturalist Vol. 39, No. 1 Table 5. Percentage composition' of beetles on 16 major study sites, 1971-73. Species of beetle Site Aph rub Chi abn Con Ian Ele his Ele Ele obs Ele sul Emb gla Gly Hyl del Sph Tri Tro tub Other^ 1 2 " 3 6 4 6 7 8 10 10 3 13 4 14 18 19 20 4 22 ■■ 23 27 28 30 35 3 3 17 10 13 4 " 23 11 16 L9 4 4 1 ° 19 20 8 ° 30 15 6 ° 3 6 8 36 5 5 27 18 5 ° ° 5 13 ' 8 5 3 5 3 6 17 (8) ° 5 ° 20 (10) ° ° ° 11 (17) 4 7 40 26 24 (2) (6) (2) 11 5 24 (7) " 4 4 24 (5) ° ° 4 11 19 (10) o o 6 5 57 (4) 45 4 15 (4) " 11 6 9 38 (3) • ° 9 69 (5) 36 3 3 25 (5) 52 22 (4) 21 15 9 (5) 'The nearest whole percentage is Hsted for those which constitute at least 3 percent of the specimens collected. However, percentage is relative to species of beetles collected. \n asterisk indicates presence in numbers less than 3 percent. 'Number in parentheses indicates number of other species present, each one less than 3 percent of the total composition. three years. Four of the 19 species that oc- curred in relative abundance in 1972 and 1973 were not present in 1971. One species present in 1971 was not taken the other two years, one was present only in 1972, and an- other only in 1973. Relative abundance of individuals differed between years. Two species were more abun- dant in 1971 than in other years, 12 were most abundant in 1972, 6 were most abun- dant in 1973, and 2 were about equally abun- dant in 1972 and 1973. In 1972, Chilometo- pon abnorme was 30 times as abundant, Embaphion glabrum 2 times as abundant, Hylocrinus delicatulus 2 times as abundant. and Sphaeriontis miiricata 4 times as abun- dant as in 1973. In 1973, Aphoditis rubidus was 24 times as abundant, Cimbocera con- spersa 4 times as abundant, and Eleodes caudifera 8 times as abundant as in 1972. In 1972 and 1973, Eleodes extricata was 11 times as abundant as in 1971. In July and August, Chilometopon ab- norme, Eleodes caudifera, Eleodes hispilabris, and Eleodes sulcipennis were most abundant in 1971. Eleodes extricata was more abundant in 1972 and 1973 than in 1971. Cononotus lanchesteri and Embaphion glabrum were about equally abundant all three years. ANTS FROM NORTHERN ARIZONA AND SOUTHERN UTAH Dorald M. Allred' and Arthur C. Cole- Abstract.— Ants of 22 species were collected in can pit-traps from 16 different vegetative associations to deter- mine distribution, seasonal and annual occurrence, and population as bases for monitoring environmental impact. Thirteen species were sufficiently abundant and distributed to qualify as indicator species. Mymiecocystus mexi- canus was the most widespread ecologically. Pogonomymiex occidentalis was the most abundant species, but second in ecological distribution. The greatest number of species was found in the juniper-£p/jgdra-grass association, and the fewest species in Ephredra-Coleogyne-Grayia. In June 1971, ecological studies were in- itiated by the Center of Health and Environ- mental Studies at Brigham Young University to establish baselines to determine the envi- ronmental impact of the Navajo Generating Station near Page, Arizona, and the proposed Kaiparowits Generating Station in Kane County in southern Utah.' Field studies of ar- thropods were conducted from July to Sep- tember in 1971, and from May to August in 1972 and 1973. Of the 12 sites operated in 1971, the 16 in 1972, and the 12 in 1973, some were studied only one year, others two, and some all three years. The trapping method, study sites, and pre- dominant vegetation on each site are de- scribed by Allred and Tanner (1971, Great Basin Nat. 39:89-96). The ants were identi- fied by Arthur C. Cole Species Commonly Collected Camponotus viciniis Mayr is common and widespread in the arid and semiarid western United States. Nesting generally beneath stones and logs, it is a strong and highly suc- cessful contender for space and food. One specimen was taken at site 1, 106 at site 2, 27 at site 6, 54 at site 19, 41 at site 27, and 5 at site 28. Largest numbers were taken at site 19, a Coleogyne community. Ants were taken from July to September in 1971, most abundantly during the latter two months; April to September in 1972, most abundantly in August; and May to August in 1973, also most abundantly in August. Comparison of July and August collections for the three years showed twice as much activity in 1972 and 1973 than in 1971. For the period of May to August, only slightly more activity was noted in 1972 than in 1973. Conomymia insana (Buckley), known pre- viously as Dorymynnex pyramiciis (Roger), has circular and semicircular crater mounds that are characteristic components of desert and semidesert regions, and the agile workers nin rapidly over the soil in great numbers during their diurnal foraging. One of the most common species of ants collected, 238 specimens were taken from 14 of the 19 sites. Largest numbers were found on site 27, a juniper-pinyon community. Ants were found in July and August of 1971, most abundantly in July; from May to September of 1972, most abundantly in August; and from May to August of 1973, also most abun- dantly in August. Comparison of July and August for the three years showed about equal numbers of ants in 1971 and 1972, but only one-fifth as many in 1973. For the peri- od of May to August, four times as many ants were taken in 1972 than in 1973. Crematogaster depilis Wheeler forms large colonies beneath stones or in nests marked by irregular craters of soil. Of the total 192 specimens taken from 11 sites, the largest numbers were found on sites 1 and 2, Ephedra-Vanclevea-grass and juni- 'Department of Zoology and Life Science Museum, Brigham Young University, Provo, Utah 84602. 'Department of Zoology and Entomology, University of Tennessee, Knoxville, Tennessee 37916. 'These studies were funded cooperatively by Salt River Project, Arizona Resources Corporation, and Southern California Edison. 97 98 Great Basin Naturalist Vol. 39, No. 1 per-Ephedra-grsLSS communities, respectively. In 1971, ants were found in July and August, more abundantly in August; in April and from June to August in 1972, most abun- dantly in July; and in May, June, and August of 1973, most abundantly in May. In July and August of the three years, twice as many ants were taken in 1971 than in 1972, and 18 times as many as in 1973. Formica ohtusopilosa Emery nests in dry, sunny areas of rather coarse soil. The large entrance is surrounded by an irregular array of pebbles. Workers forage rapidly over the insolated soil surface. Forty-one specimens were taken at site 3, an Ep/iedra-grass community, and 18 at site 28, an Artemisia-grass community. The ap- parent absence of this species in other areas is unusual. Ants were taken in 1971 in July and August, predominantly in July; in 1972 from June to September in about equal num- bers each month; and in about equal numbers from May to August in 1973. Activity for July and August in 1971 was about equal to the same months of 1972, but about five times greater than in 1973. For the period of May to August, four times as many ants were taken in 1972 than in 1973. Formica rufibarbis gnava Buckley forms rather large colonies in earthen nests general- ly marked by an irregular crater or mound. Seventy-one specimens were taken at site 2, a ixmiper-Ephedra-grsiSS community, and 7 specimens at site 3, an Ephedra-grass habitat. All but one of the specimens were taken in 1971, mostly during July. Only one specimen was taken in 1973, in May. Iridomymiex pruinosum (E. Andre) is a successful and common component of the desert and semidesert biota. Although it often constructs small circular or irregular craters of fine soil particles, it frequently nests be- neath stones and debris. Workers forage rap- idly in files even when the soil surface has a high temperature. Ants were taken from eight sites. More were taken from site 6, an Artemisia commu- nity, than from any other site. They were found only in August of 1971, and from June to August in 1972. Monomorium minimum (Buckley) is a min- ute ant that nests beneath stones and detritus, or in open soil with or without a crater. It forms populous colonies. Thirty-five specimens were taken from seven sites. Largest numbers were found at site 28, an Artemisia -grass community. Ants were taken in August 1971, from May to Sep- tember in 1972, and only in May of 1973. Most of the specimens in 1972 were taken in July. Thirty-one of the specimens were taken during 1972. Myrmecocystus mexicanus Wesmael builds nests in unshaded soil. Each nest is usually surmounted by a mound bearing a single, large circular entrance of firmly packed coarse sand. The workers are nocturnal for- agers. Ants of this species were some of the most abundant and the most widespread of all those found. A total of 652 was taken from all 19 sites, the numbers not different signifi- cantly from one site to another. They were found during each of the summer months that studies were made. Largest numbers were found in August of 1971, in July and August of 1972, and in May and June of 1973. Comparison of July and August for each of the three years showed about equal numbers in 1971 and 1972, but only one- third as many in 1973. For the period of May to August, half again as many ants were taken in 1972 as in 1973. Myrmecocystus melliger Forel constructs its nests in rather loose soil marked by a cir- cular or semicircular crater. These ants run rapidly over the ground during their diurnal foraging, and are a conspicuous though famil- iar element of desert and semidesert areas. Seventy-one specimens were taken from 12 sites. No significant difference was noted be- tween the numbers of ants at each of the sites. They were taken in July and August of 1971 in about equal numbers; from June to August of 1972, predominantly in July; and in small numbers in May, July, and August of 1973. Comparison of July and August among the three years showed about equal numbers in 1971 and 1972, but only one-fifth as many in 1973. For the period of May to August, four times as many ants were taken in 1972 as in 1973. Pheidole bicarinata paiute Gregg is a small ant with dimorphic workers that generally nests beneath stones in sand or fine soil. Both March 1979 Allred, Cole: Ecology of Ants majors and minors forage largely for seeds during the cooler daytime hours. A total of 198 specimens was taken from 12 sites. Numbers were highest at sites 6 and 28, which were Artemisia communities. In 1971, ants were taken from July to Septem- ber, mostly in August; from May to Septem- ber in 1972, predominantly in September; and only in June and July of 1973, pre- dominantly in June. Comparison of July and August for the three years showed twice as many in 1971 as in 1972, and 25 times as many as in 1973. For the period of May to August twice as many were taken in 1972 as in 1973. Pheidole sitarches soritis Wheeler is anoth- er dimorphic harvester that constructs small, circular crater nests or sometimes lives be- neath stones in unshaded areas. A total of 110 specimens was taken from 10 sites. Largest numbers were found at site 3, an Ephedra-grsLSS community. In 1971 they were found in July and August, mostly July; from June to August in 1972, predominantly in August, and from June to August in 1973, predominantly in June. Comparison of July and August for the three years showed only two-thirds as many in 1972 as in 1971, and only one-fourth as many in 1973. For the pe- riod of May to August, numbers were about equal in 1972 and 1973. Pogonomyrmex occidentalis (Cresson) is a common harvester whose numerous, large, conical or subconical mounds, each sur- rounded by an area cleared of plants, are some of the most conspicuous features of the desert. The nests are usually in coarse, gravelly soil, and both the underground por- tion and the superstructure contain chambers and galleries in which brood is raised and seeds are stored. Ants of this species were the most abun- dant but not the most widespread of all taken. A total of 800 specimens was taken from 15 of the 19 sites. Ants of the species Myrmecocystus mexicaniis were more wide- spread than P. occidentalis, but not as abun- dant. This species was most abundant on site 23, although sites 8, 13, and 14 also had rela- tively high populations. They were found during each of the summer months that field studies were made. In 1971 and 1972 largest numbers were found in August, and in 1973 in June. Comparison of July and August showed four times as many in 1972 as in 1971, and 13 times as many as in 1973. For the period of May to August, eight times as many were taken in 1972 as in 1973. Pogonomyrmex rugosus Emery, another large harvester species, constructs nests sur- mounted by a low gravel mound or disc with a large, irregular central entrance. The work- ers vigorously defend their nests and sting readily. Thirty-three specimens were taken from five sites. Largest numbers were found at site 20, a grass community. Only one was taken in 1971 in July, and the balance in about equal numbers from July to September in 1972. Species Rarely Collected Camponotus semitestacea Emery: one speci- men 6 June 1972, 3 on 6 July 1972, 3 on 5 August 1972, one on 30 April 1973, site 2— juniper-Ep^edra-grass; 3 on 4 July 1973, site 6— Artemisia. Conomyrma bicolor (Wheeler): 5 specimens 15 August 1972, site 19— Coleogyne. Crematogaster mormonum Emery: 7 speci- mens on 18, 19 August 1971, site 9-At- riplex Artemisia. Liometopum luctuoswn Wheeler: 2 speci- mens 19 June 1972, site 28— Artemisia grass; 12 on 12 May 1973; 15 on 1 July 1973, site 27— juniper-pinyon Myrmecocystus mimicus Wheeler: 7 speci- mens on 14, 16 August 1971, site 2— ]uniper-£p/iedra-grass; one on 6 June 1972, site S—Ephedra-grass; one on 14 August 1972, site 22-Coleogyne. Pogonomyrmex imberbiculus Wheeler: 3 specimens on 18, 20 August 1971, 2 on 11 August 1972, site 9-Atriplex-Arte- misia. Pogonomyrmex suhnitidus Emery: 4 speci- mens 18-20 August 1971, one on 10 April 1972, 2 on 12 July 1972, 3 on 10, 12 August 1972, one on 11 September 1972, site 10— £p/iedra-grass. Veromessor lobognathus (Andrews): one spec- imen 19 August 1971, 2 on 10, 11 Au- gust 1972, 2 on 11 September 1972, 6 on 6, 7 June 1973, site lO-Ephedra- grass; one on 7 July 1972, site 4— Co- leogyne; 3 on 12 July 1972, site 8- Gra j/ia-grass. 100 Great Basin Naturalist Vol. 39, No. 1 Discussion These studies were established in major vegetative types within a 48-km perimeter of the proposed sites of the electric generating stations. Retention of study sites beyond the first year was based on vegetative type, di- rection from the potential source of pollu- tion, and especially the species and relative abundance of organisms present that could be used as indicator species to monitor environ- mental changes. Some sites were dis- continued after one season because of in- accessibility, discontinuous vegetative analysis, and no specific climatic and edaphic data. In order to compare populations and sea- sonal changes, the numbers of ants collected were adjusted to the number of trapping at- tempts. The normal variability in seasonal and annual populations, slightly different trapping periods within the same month, and the influence of periodic and abrupt climatic changes on the activity of the ants during a given trapping period were ignored. Pit traps are effective primarily for ground-dwelling arthropods that move on the ground more frequently than they fly. The traps involve minimum effort and time, can be used effectively for those species which may be so trapped, and adequately determine relative abundance and distribution within the limitations of time, economy, and logis- tics. The number of trapping attempts is shown in Table 1. Thirteen of the 22 species found in this study are sufficiently abundant and dis- tributed that they may be used as indicator species to determine environmental changes (Table 2). In this study relative numbers of individ- uals are referred to as "activity," inasmuch as populations were measured only by above- ground activity and not numbers of colonies or individuals within those colonies. An as- sumption is made that above-ground activity and abundance is directly proportional to the number of colonies and individuals. Greatest activity of the ants occurred in July and August of 1971 and 1972, and in June of 1973 (Table 3). The total number of species varied only slightly during the three years (Table 4). Some study sites showed significant varia- tions where data were available for all three years. On 6 of 8 such sites, a decrease in the number of species was noted in 1973. Where only two years' data were available for 5 sites, one site increased in number of species in 1972 over 1971, 3 decreased in 1973 over 1972, and one was equal for 1972 and 1973. Nine of 12 sites showed a species decrease in 1973 over one or both the two previous years. Table 5 shows the variety of predominant ants for each of the study sites. Myrmeco- Table 1. Number of trap-days' for pit traps on 16 major study sites, 1971-1973. Site 1971 1972 1973 Total 1 60 120 90 270 2 60 180 90 330 3 60 180 90 330 4 60 120 ° 180 6 60 90 90 240 8 60 90 90 240 10 60 120 90 240 13 60 150 90 300 14 60 150 90 300 19 90 ° 90 20 90 ° 90 22 150 ° 150 23 120 90 210 ' 27 120 70 190 28 120 70 190 30 150 90 240 Total 510 2,040 1,040 3,590 'Number of traps multiplied by number of days operated. "Not operated during year indicated. Table 2. Numbers and distribution of ants' on 16 major study sites, 1971-1973. Total No. sites number where Species taken found Pogonoinymiex occidentalis 800 14 Mynnecocystus mexicanus 652 16 Conomymia insana 2,38 12 Caniponotus vicintis 234 6 Pheidole paitite 198 10 Crematogaster depilis 192 10 Phiidole sitarches 110 9 I'onniva rtifiharbis 78 2 Mynnecocystus melliger 71 11 Formica obtiisopilosa 61 2 Monomorium minimum 35 7 Pogonomijnnex rugosus 33 4 Iridomynnex pruinosum 26 7 'Only those species are included of which more than ; taken. March 1979 Allred, Cole: Ecology of Ants 101 Table 3. Total number of ants of all species collected in pit traps at 16 major study sites, 1971-1973. No. ants collected Month 1971 1972 1973 Actual Adjusted' Actual Adjusted' Actual Adjusted' April May June ; 34 46 266 54 69 319 116 111 151 266 July 184 736 649 649 88 176 August September 424 27 721 216 701 96 841 307 76 152 'Numbers adjusted to number of trap days. 'Traps not operated. Table 4. Number of species of ants captured in pit traps on 16 major study sites, 1971-1973. Site Predominant vegetation 1971 1972 1973 Total 1 Ephedra-Vanclevea-Grass 2 junipeT-Ephedra-Grass 3 Ephedra-Grass 4 Coleogyne 6 Artemisia 8 Grayia-Grass 10 Ephedra-Grass 13 Grayia-Ephedra-Grass 14 Coleogyne-Grayia-Ephedra-Gi 19 Coleogyne 20 Grass 22 Coleogyne 23 Ephedra-Coleogyne-Grayia 27 Juniper-Pinyon 28 Ar/emisia-Grass 30 Ephedra-Grass 7 8 8 10 10 9 10 13 7 9 4 11 4 7 ° 7 7 6 3 8 5 5 3 6 5 8 4 8 5 7 4 7 7 4 7 9 • 9 5 • 5 7 ° 7 3 3 3 8 6 9 8 4 9 6 2 6 Total 20 20 'Not trapped during year indicated. Table 5. Percentage composition' of ants on 16 major study sites, 1971-1973. Species of ant Cam Con Cre For For Mon Myr Myr Phe Phe Pog Pog Site vie ins dep obt gna min mex pla pai sor occ rug Other^ 1 » • 26 ° 28 » 18 ' • (1) 2 26 • 12 18 20 ° ' ' ° (3) 3 14 16 " ° 10 ° 14 20 17 ° (1) 4 - " 41 ° ' 38 (1) 6 19 26 ° 36 ° (3) 8 " • 23 64 (1) 10 51 ° ° 22 (3) 13 ° 14 ° ° 69 (1) 14 ° 24 " ° 63 (1) 19 55 " " 16 ' ° ° 20 27 ° ° ° 49 22 53 ° 27 ° * (1) 23 » 27 72 27 15 34 24 ° ° 11 (3) 28 » 15 16 19 ° 28 ° ° (1) 30 21 ° ° 29 ° 43 'The nearest whole percentage is hsted for those which constitute at least 10 percent of the specimens collected. However, percentage is relative to all species of ants collected. An asterisk indicates presence in numbers less than 10 percent. -Number in parentheses indicates number of other species present, each one less than 10 percent of the total composition. 102 Great Basin Naturalist Vol. 39, No. 1 cystus mexicanus was on all 16 of the sites evaluated, but not significantly abundant on 2 sites. Pogononiyrmex occidentalis was on 14 of the sites, but of significant composition on only 9. Only 2 sites had the same species composition (sites 8 and 30), but the relative percentages of each species differed between them. Comparison of sites which were most alike in predominant plant species showed some significant differences. Coleogyne sites 4, 19, and 22 had only one species of ant in com- mon. Site 4 had three unique species, and site 22 had two unique species. Comparison of sites 14 and 23, which also contained signifi- cant amounts of Coleogyne, showed both sites with one species common to each and to the other Coleogyne sites, both sites with two species common to each and to site 4, and site 14 with one species common to site 22. Comparison of £p/iedra- grassland sites 3, 10, and 30 showed only one species common to all three. One species was common to sites 3 and 30, one species to 10 and 30, three spe- cies unique to site 3, two to site 10, and one to site 30. Comparison of sites 1 and 20, which also had significant amounts of Eph- edra and grass, showed one species common to each and to sites 10 and 30, one species common to each and to site 3, one common to each and site 30, site 1 with one species common to sites 3 and 30, and with each site with one unique species. Artemisia sites 6 and 28 had only one spe- cies in common. Three species were unique to site 6, and four species to site 28. Grai/ia-grass sites 8 and 13 had three spe- cies in common, and each had one unique species. Juniper woodland sites 2 and 27 had three species in common, site 2 had two unique species, and site 27 one unique species. Annual differences in species composition varied between the three years. Two species that were active in 1972 and 1973 were not taken in 1971. One species active in 1973 was not taken in the other years, and one species present in 1972 was not taken in the other years. Three species present in 1971 were not taken in 1972 or 1973. One species taken in 1971 and 1973 was not found in 1972, and five species taken in 1971 and 1972 were not taken in 1973. Relative activity (measured by the number of specimens caught in traps) of individual species differed between years. One species was more active in 1971 than in other years, 10 were most active in 1972, 2 in 1973, and one more active in 1972 and 1973 than in 1971. In years when comparisons could be made for similar months, most species were much more active in 1972 than in 1973. Camponotus vicinus was only slightly more active; Conomyrma insana, Formica obtu- sopilosa, and Myrmecocystus melliger were 4 times as active; Crematogaster depilis 3 times as active; Pheidole paiiite 2 times as active; Myrmecocystus mexicanus half again as ac- tive; Pogononiyrmex occidentalis 8 times as active; and Monomorium minimum 33 times as active in 1972 as in 1973. Pheidole soritis was about equally active in 1972 and 1973, and Veromessor lohognathus was 2 times as active in 1973 as in 1972. Comparison of the months July and August for the three years showed that Crematogas- ter depilis, Pheidole paiute, and Pheidole so- ritis were most active in 1971; Iridomyrmex pruinosum, Monomorium minimum, Pogono- niyrmex occidentalis, P. rugosus, and Vero- messor lohognathus in 1972; Conomyrma in- sana, Formica obtusopilosa, Myrmecocystus mexicanus, M. placodops, and Pogononiyrmex subnitidus in 1971 and 1972; and Camp- onotus vicinus in 1972 and 1973. TABLE OF CONTENTS Phlox longifolia Nutt. (Polemoniaceae) complex of North America. Frederick J. Peabody 1 Diatom floristics and succession in a peat bog near Lily Lake, Summit County, Utah. Shobha A. Jatkar, Samuel R. Rushforth, and Jack D. Brotherson 15 Vegetational response to three environmental gradients in the salt playa near Goshen, Utah County, Utah. Michael G. Skougard and Jack D. Brotherson 44 Distribution of sculpins in the Clearwater River Basin, Idaho. O. Eugene Maughan and Gary E. Saul 59 Phytoseiid predators of mite pests in Utah apple orchards. Clive D. Jorgensen and Vichitra Mongkolprasith 63 Subspecies specificity of gall forms on Chrysothainnus nauseosus. E. Durant McArthur, Charles F. Tiernan, and Bruce L. Welch 81 Homing by a pigmy rabbit. Jeffrey S. Green and Jerran T. Flinders 88 Beetles from the environs of Lake Powell in southern Utah and northern Arizona. Dorald M. Allred and Vasco M. Tanner 89 Ants from northern Arizona and southern Utah. Dorald M. Allred and Arthur C. Cole 97 NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be di- rected to Brigham Young University, Stephen L. Wood, Editor, Great Basin Xataralist, Provo, Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be sim- ilarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. 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 Edi- tors Style Manual, Third Edition (AIBS 1972) in preparing the manuscript. An abstract, about 3 percent as long as the text, but not exceeding 200 words, written in accordance with Biological Abstracts guidelines, should precede the introductory paragraph of each ar- ticle. Authors may recommend one or two reviewers for their article. All manuscripts re- ceive a critical peer review by specialists in the subject area of the manuscript under con- sideration. 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Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul- tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In gener- al, only previously unpublished manuscripts of less than 100 printed pages in length and per- taining to the biological and natural history of western North America are accepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natu- ral history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manuscripts are subject to the approval of the editor. Subscriptions. 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All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back 8-79 650 40666 ISSN 0017-3614 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 39 June 30, 1979 No. 2 REVIEW OF TULAREMIA IN UTAH AND THE GREAT BASIN Harold E. Stark' Abstract.— This is a compilation of tularemia research conducted in Utah, particularly at U.S. Army Dugway Proving Ground (DPG), Utah, and an evaluation of this information in relation to the current status of tularemia studies. A brief history of tularemia in Utah and a review of field and laboratory studies are included. Human cases of tularemia occur throughout Utah during all seasons of the year. An analysis of recent human disease reveals a concentration of cases in rural areas, with a greater seasonal occurrence in late summer and early fall. Research on tularemia as a zoonotic in- fection in and around the U.S. Army Dugway Proving Ground (DPG), Utah, has established the existence of natural foci of infection, cy- cles of activity, and probable reservoir hosts and vectors. In addition, studies have been directed toward determination of survival times of the organism as aerosols and as con- taminants on surfaces in the laboratory and in nature under varying field conditions. Field and laboratory work also have been con- ducted at Brigham Young University and the University of Utah; several species of deer fly (Chrysops spp.) were found infected in na- ture. Human tularemia is commonly a rural dis- ease, probably with an eight-year cyclic ten- dency in Utah. It is transmissible to man through direct contact with the host, con- tamination of water and food, vectors, in- halation of dust as from tick feces while shearing sheep, and, uniquely in the Great Basin, by the bite of deer flies. There have been several review articles on tularemia in recent years; Jellison and Parker (1945), Bell (1965), Olsuf'yev and Rodnev (I960), Hopla (1974), and Olsen (1975) are examples. In addition, several bibliographies have been compiled, e.g.: the U.S. Army Chemical Corps (1958), Hoogstraal et al. (1970-1972), and Pollitzer (1967). Cox (1964), at Brigham Young University, prepared a "Bibliography of Tularemia" with references arranged by subject matter, similar to the one published by the U.S. Army Chemical Corps. Jellison was author (1950) and coauthor (1945, 1951, 1956), of specific and general ar- ticles on tularemia (many of them on tula- remia in Utah), and he prepared a current re- view and bibliography on tularemia (1974). Much of the information given here is a re- view of large quantities of data from DPG re- ports and records not ordinarily available to the scientific community. Since the data are voluminous, not all can be analyzed; many DPG reports and articles are referenced for further analysis (if desired) by the reader. Other portions of this work constitute a liter- ature survey. 'Environmental and Ecology Branch, U.S. Army Dugway Proving Ground, Dugway, Utah ! 103 104 Great Basin Naturalist Vol. 39, No. 2 History The etiologic agent of tularemia, Franci- sella tularensis, was not identified until this century (McCoy 1911, McCoy and Chapin 1912). However, tularemia may have ap- pared earlier among humans in rural Utah. Pearse (1911), a Brigham City physician, studied six human cases of an imknown dis- ease which occurred in August 1908 in the Brigham City-Tremonton area. It is believed to be the first clinical description of tula- remia in the English language. Pearse left the disease unnamed. Later medical accounts re- viewing epidemiologic and clinical aspects of the disease assumed deer fly bites were asso- ciated with it, although no infected insects were captured or identified. Francis, another physician, first isolated F. tularensis from wild mammals and their ecto- parasites in 1919 and conducted detailed studies near Holden in the Delta area of Mil- lard County (Francis 1921, 1927). Before 1920 he referred to the disease as "Pahvant Valley plague" and "deer fly fever." Later (1921) he established and used the name tula- remia for the disease. Simpson (1929) wrote a textbook on medical aspects of the disease, assembling knowledge to that time. He asso- ciated the cases in Utah largely with deer fly bites. Etiologic Agent Francisella is distinctive and only distantly related to other bacterial organisms. Buchan- an and Gibbons (1974) gave no hierarchical arrangement for it. The genus is placed with Brucella, to which it is not closely related," in a group of genera of uncertain affiliation en- titled "Gram-negative aerobic rods and coc- ci." There has been no general compilation of strains published to date, although there are significant differences in levels of viru- lence among strains of F. tularensis. Yet viru- lence alone does not provide a basis for clas- sification, because under laboratory culture and storage it can change. Different labora- tory hosts express different levels of resist- ance and display varying pathogenesis. Bio- chemical reactions (glycerine fermentation and the presence or absence of citruUine ureidase) have been related to virulence (Marchette and Nicholes 1961). Some strains resist streptomycin. Eigelsbach, Braun, and Herring (1951), Skrodsky (1966), and Domin- owska (1967) observed a correlation between the colonial morphology and the pathogeni- city and immunogenic properties of a given isolate. Based on virulence, chemical reactions, morphology, geographic origin, epizootiol- ogy, epidemiology, vectors, reservoirs, associ- ation of different habitats, and modes of transmission, two basic subspecies of tula- remia have gained recognition. These are commonly referred to as types A and B (Bell 1965). Subspecific designations also have been made referring to types A and B. The less virulent of the two is palaearctica (Type B) and is more frequently associated with ag- ricultural areas and lotic waters, and may be ' maintained by chronic tularemia nephritis in muskrats, beavers, and voles. Voles may be the primary reservoir in some areas in west- ern North America. Chronically infected ver- tebrates urinate onto watersheds, sometimes causing widespread and protracted human epidemics (Bell and Stewart 1975). Human infection with palaearctica has oc- curred during threshing operations in the USSR. Possible similar occurrence of human tularemia in the Great Basin is discussed later imder tularemia in soils. Another proposed name, holarctica, for Type B, implies that this type occurrs throughout all land masses of the Northern Hemisphere. Though holarc- tica is less restrictive in concept, palaearctica has taxonomic priority. Francisella t. tula- rensis (Type A, also designated F. t. nearc- tica) is more virulent and is frequently associ- ated with infection in lagomorphs (hares and rabbits) and nonaquatic (xeric or mesic) ro- dents, more frequently involves human cases contracted during hunting, and is often asso- ciated with vectorborne transmission than is palaearctica (Bell 1965). Besides the currently recognized sub- 'There is some serologic cross reaction between F. tularensis and Brucella (Hopla 1974, Quan 1978). June 1979 Stark: Review of Utah Tularemia 105 species, two additional nomenclatural desig- nations have been proposed, }aponica (Ro- dianova 1967) (for Japanese isolates) and mediasiatica (Aikimbaev 1966) (for a central Asian strain). Aikimbaev (1966) and Olsul'yev (1970) regard the latter as primitive. Aikim- baev proposed subspecific status for media- siatica. Taxonomic status has not been eval- uated for either of these two names. The tularemia organism in nature is ubiq- uitous, but simultaneously demonstrates ubiety with regard to strain differentiation. Numerous strains have been recognized. These often have been designated with num- bers or initials. At DPG, nine strains are maintained and studied: Jap 4, Ohara, Live Vaccine, S. C, Russ, Max, 38, 38A, and Schu 5. Similar groups have been kept at several educational institutions, the U.S. Public Health Service, Rocky Mountain Laboratory at Hamilton, Montana; Centers for Disease Control at Fort Collins, Colorado, and At- lanta, Georgia; and at several state health de- partments. Several strains may occur in one locality (e.g.. Prince George County, Vir- ginia, where several strains came from a single species of tick— Haemaphy sails lepo- rispalustris). Karlsson et al. (1970) and Karls- son and Soderlind (1973) obtained 50 strains from a limited area in Sweden; most of these came from a single species of tick. Many publications list strains collected locally from nature or developed in the laboratory. In Stoenner et al. (1959) and Marchette and Nicholes (1961), the strains DPG-1,2,3,4,5,6, SKV- 1,2,3, and 9-K-161 are reported as iso- lates from ticks {Dermacentor parumpertus, Hemaphysalis leporispalustris, and others), cottontails {Sylvilagus audobonii), and jack- rabbits (Lepus californicus) around DPG. Marchette et al. (1961) listed 18 cultures kept at DPG and at the University of Utah, Salt Lake City. A staff report on tularemia at DPG (No. 88U; USA-DPG 1962b) listed 26 strains (18 from DPG) used in susceptibility and vector studies of transmission. There is no record of preservation of these isolates. Some strains are preserved in adjacent states. Dr. Thomas J. Quan of the Plague Branch of the Communicable Disease Center, Fort Col- lins, Colorado, has strains SKV-2 and DPG-1 as part of his F. tularensis collection. Mr. Scott Stewart, Rocky Mountain Laboratory, Hamilton, Montana, preserves numerous strains by lyophylization, but effort is not de- voted to identification and cataloging; it is more convenient to obtain fresh material from nature for ongoing research. Storage and preservation or cultivation (es- pecially in egg yolk) in the laboratory reduc- es virulence of the organism (Owen 1970). Green (1943) increased virulence of unnamed strains of F. tularensis by passage through cottontail rabbits and hares. On the other hand, he contended that passage through grouse reduced virulence. Owen et al. (1961) failed to enhance the virulence of F. tula- rensis by experimentally passing strains through hosts or ectoparasites. Some strains have considerable vitality for sustaining their characteristics. There are great variations of virulence in strains isolated in nature. Older literature on virulence refers to resistance, susceptibility, and mortality in human in- fections. No assignment of strains, even as far as types A or B, is possible for these old isola- tions. No classification or cataloging of F. tularensis strains appears possible at the pres- ent time. Buchanan and Gibbons (1974) sug- gested a nomenclature that has had partial acceptance at best. Thus, resolving the status of the number and identity of the strains of F. tularensis (other than the two principal types) remains an unresolved problem. Research on Tularemia in Utah AND THE Great Basin, 1945-1975 Table 1 summarizes research on tularemia in nature in the Great Basin. These in- vestigations are discussed in the following section. Krinsky (1976) devoted about half of his review of tularemia in tabanid flies (Chry- sops spp.) to the historical occurrence of tula- remia in nature in Utah and to research pub- lished by Utah-based authors. Krinsky's high regard for studies of tularemia in Utah prompted an expanded review. Distribution, reservoirs, and cycles in nature.— The occurrence of tularemia in Utah differs from that in other parts of the world. Rodenwaldt et al. (1952) believed F. tularensis to be ubiquitous in the Holarctic Realm and to occur in permanent and well- defined epicenters where it is maintained, and from which it may spread. The area from 106 Great Basin Naturalist Vol. 39, No. 2 Table 1. Summary of basic research on tularemia in the Great Basin, 1945 to 1975. Principal Investigators' Date Institution^ Subject of research Jellison, Parker Jellison Jellison, Kohls, Philip Jellison, Kohls Rodenwaldt Woodbury, Parker Woodbury U of U staff Philip, Bell, Larsen Allred, Stagg, Lavender Stagg, Tanner, Lavender Parker, Johnson Vest, Marchette Stoenner, Holdenried U of U staff Parker, Olsen, Dolana E&E staff Lundgren, Marchette, Nicholes ditto Gebhart, Thorpe 1945 1950 1951 1956 1952 RML RML RML RML U.S. Navy 1953 UofU USA-DPG 1954 UofU USA-DPG 1955 USA-DPG 1955 RML 1956 USA-DPG 1956 UofU USA-DPG 1957 UofU USA-DPG 1958 UofU USA-DPG 1959 RML USA-DPG 1961 thru 1969 UofU USA-DPG 1970 thru 1972 EcoDynamics USA-DTC 1973 to present USA-DPG 1961 UofU USA-DPG 1962 UofU USA-DPG 1962 UofU USA-DPG Rodents, rabbits, and tularemia; Sylvilagus ssp.— basic reservoir and source of human tularemia Deer fly distribution Tularemia in sheep Tularemia in muskrats and humans around Utah Lake Concept of epicenters Special report on tularemia Tularemia and biotic communities Ecology of tularemia transmission Tularemia in jackrabbits in Nevada Transmission by Dermacentor parwnapertus Experimental infection of native animals Attempted transmission by fleas Transmission from infected carcasses Isolates of tularemia from Great Salt Lake Desert area Annual reports of surveillance Annual reports of surveillance Annual reports of surveillance Immunity of host; virulence of pathogen Cutaneous allergic reaction Review of literature June 1979 Stark: Review of Utah Tulahemia 107 Arkansas to southern Illinois constitutes one major epicenter in North America. Another well-defined epicenter exists in the western part of the continent, of which Utah (espe- cially the Delta area) is a part. Later pub- lications attach no particular significance to these epicenters of tularemia as defined by Rodenwaldt et al. Maximov (1960), however, regarded tularemia as an important natural regulator of rodent hosts and reported that it occurs in favorable foci in the USSR as re- lated by the concept of "landscape epide- miology." Jellison and Parker (1945) proposed that cottontails (Sylvilagus spp.) are basic reser- voir hosts responsible for 90 percent of hu- man tularemia in North America. Several species of deer fly are the principal vectors to man from zoonotic sources (possi- bly infected hares) in Utah. Throughout the remainder of the world, ticks and sometimes mosquitoes (Rodenwaldt et al. 1952) are the principal vectors to man. Other sources of human tularemia infection include contact with the infected hosts, contaminated water, or aerosols. Tularemia appears to occur widely in a va- riety of environments and hosts. Few areas are as ecologically diversified as Utah. Tula- remia is widespread from the Great Salt Lake Desert to the Uinta Mountains. It has trans- mission cycles involving waterborne tula- remia, as with muskrat trappers around Utah Lake (Jellison, Kohls, and Philip 1951); air- Thorpe, Marcus, Sidwell 1962 thru 1967 UofU USA-DPG Cabelli etal. 1964 USA-DPG Cox 1964 BYU Cox 1965 BYU Thorpe et al. (1965) 1951 thru 1964 UofU USA-DPG Thome 1966 USA-DPG Johnson 1966 UofU USA-DPG Vest et al. 1965 UofU USA-DPG Knudsen, Rees, Collett 1968 UofU USA-DPG Klock, Fukushima, Olsen 1973 UDH USA-DTG Olsen 1975 EcoDynamics USA-DTC Phagocytosis and other cellular factors relating to resistance Transmission by birds Bibliography of tularemia Tularemia in deer flies Tularemia in wildlife and livestock in Great Salt Lake Desert Tularemia in soils at DPG Ticks of DPG Five-year review of tularemia field infections in western Utah Tularemia in deer flies in Salt Lake County; trapping tabanids Tularemia epidemic: Delta, Grantsville Review of tularemia 'See bibliography for complete citation. 'Institution abbreviations; BYU = Brigham Young University, Provo, Utah DPG = U.S. Army Dugway Proving Ground, Utah DTC = Deserel Test Center, Fort Douglas, Utah E&E = Epizoology and Ecology Branch prior to 1972; now Environmental and Ecology Branch, DPG RML = Rocky Mountain Laboratory, Hamilton, Montana UDH = Utah State Division of Health USA = U.S. Army U of U = University of Utah, Salt Lake City, Utah 108 Great Basin Naturalist Vol. 39, No. 2 borne tick feces as with sheep shearers (Jelli- son and Kohls 1956); direct contact, as with rabbit hunters; and deer fly bites as with 26 of 170 laborers at Locomotive Springs, Utah (Burnette 1936, Hillman and Morgan 1937). These latter authors reported that laborers removed their shirts while working, that sick jackrabbits were seen in the area, and 13 men were known to have killed jackrabbits or handled dead ones. The locations of initial ul- cers of the shoulders and backs of the work- men suggested, however, that jackrabbits were not directly involved as a source of hu- man tularemia. Francisella novicida was isolated and de- scribed from Ogden Bay, on the eastern side of the Great Salt Lake (Owen 1974), and has never been reported elsewhere. Thus Utah differs from all other areas of the world by harboring all three recognized taxa of Franci- sella. Epizootiology of tularemia at DPG.— In 1951 a program was begun to study and analyze plants and animals at DPG and adja- cent areas (Woodbury 1956, 1964). Emphases were placed on enumerating enzootic path- ogens as well as potential introduction of oth- er pathogens to determine possible vectors and routes of natural dissemination. Most re- search on tularemia at DPG was conducted under contract. Reports entitled "A Study of the Ecology and Epizoology of the Native Fauna of the Great Salt Lake Desert" (An- nual Summaries 1951-1969) were published by the University of Utah (USA-DPG 1951-1969). "Ecology Studies of Western Utah" 1971-1973 were prepared by EcoDy- namics Incorporated (Desert Test Center USA-DTC 1971-1973). From 1969 through 1964, special summary reports on tularemia were published as DPG in-house reports. These voluminous reports contain much raw data in tables and graphs. Additional tabula- tion, study, and synthesis of data are not within the scope of this review. The follow- ing provides a cursory summary of some re- sults. From 1951 through 1979 evidence of tula- remia existed in virtually all 46 areas of Utah in which specimens were collected and from 35 areas sampled routinely. Serologic evi- dence of the organism was found infrequently from rodents, usually in less than 1 percent of the specimens. In contrast, high rates were found for carnivores and livestock, usually greater than 11 percent, while moderate rates were recorded for deer and wild horses. Greater risk of exposure, longer lives, innate resistance, and persistence of antibodies may account for higher occurrence in these mam- mals as opposed to rodents. The incidence of antibodies was usually similar in widely dif- ferent groups of hosts in the same year. For example, in 1965 22 percent of 651 cattle samples were positive, 42 percent of 1,579 sheep samples were positive, and 25 percent of 31 carnivore samples were positive. In 1968 the proportions of samples with anti- bodies to F. tularensis were small: 0.03 per- cent for rodents, 2 percent for livestock, and 3 percent for carnivores. In other years carni- vores persisted with high rates, and rates in other animals were low. For example, in 1970 antibody rates were 4 percent in sheep, 0.4 percent in cattle, 0 percent for rodents, and 17 percent for carnivores. Antibody re- coveries from rodents and jackrabbits re- mained consistently small however, sugges- ting that rodents play a minor role in maintenance. Also of little significance to the maintenance of F. tularensis in nature is the jackrabbit, which is very susceptible and suc- cumbs promptly. This observation may ex- plain why few jackrabbits in nature possess antibodies to tularemia. A special summary report prepared by the Ecology and Epizoology research staff, Uni- versity of Utah, summarized DPG studies on host relationships for ticks, lice, and fleas (USA-DPG 1962b). The report discussed ex- perimental vector transmission studies that included immunization and susceptibility ex- periments using several strains of F. tula- rensis on laboratory mice, rabbits, guinea pigs, and deer mice. Isolations from wild mammals, birds, and livestock were tabu- lated. Ecological observations on biotic com- munities and population dynamics, particu- larly for the jackrabbit, were summarized. Serologic methods for detecting antibody ac- tivity, especially agglutinating, complement- fixing tests, and cutaneous responses were re- viewed. Some of the information in this re- port has not appeared in published literature. Stagg, Tanner, and Lavender (1956) pro- duced experimental infection of F. tularensis June 1979 Stark: Review of Utah Tularemia 109 in jackrabbits, seven species of rodents, and coyotes (Canis latrans). They found that coy- ote pups were less susceptible to tularemia than rodents. Infection of rodents via con- taminated food was difficult. When infection of rodents and jackrabbits was successful (as from aerosols), the disease usually progressed rapidly to death (3-5 days). Gebhardt and Thorpe (1962) presented a tabular review of worldwide vectors and hosts. The pathogen, vectors, hosts, epide- miology, epizootiology, and control were giv- en a generalized interpretation. Thorpe et al. (1965) reported on tularemia studies from their inception at DPG (1951 through 1964). Areas of study included the Bonneville Basin of western Utah and adjacent portions of Ne- vada, where 35 areas were sampled semi- annually or quarterly. During the 13-year pe- riod 52 isolations of F. tularensis were made, approximately half from wild mammal tissues and half from ectoparasites. The authors also tabulated information on human tularemia from 1941 through 1964. This information, updated with data from the Utah State Divi- sion of Health, is presented in this article as Figure 1 and is discussed later in the section: "Distribution of Recent Cases of Tularemia in Utah." Isolation data with respect to live- stock, wild mammals, birds, and their ecto- parasites and collections of positive sera were also tabulated. Six local epizootics were ob- served during the 13-year period. Tissue iso- lates were made from jackrabbits, which were collected in great abundance, one cot- tontail, and four rodents. Serum agglutinins were found in birds, wild mammals (includ- ing deer), and livestock (roughly 25 to 30 percent of the serum samples tested). Most isolates possessed maximal virulence; how- ever, two isolates from the cottontail and a Great Basin pocket mouse (Perognathus par- vus) were of low virulence. Thorpe et al. in- dicated that strains from North American nonaquatic hosts and ectoparasites are some- times of lower virulence. Vest et al. (1965), in a five-year study of enzootic diseases in Utah, reported that 10 strains of F. tularensis were isolated from la- gomorphs and rodents. All strains were of maximum virulence. One viable organism sufficed for a lethal dose in laboratory rab- bits, mice, hamsters, and guinea pigs. Olsen and Dolana ([EcoDynamics, Inc.] USA-DTC 1971-1973) provided additional support to Stagg, Tanner, and Lavender (1956), March- ette et al. (1961), and Thorpe et al. (1965). Olsen and Dolana found that (1) F. tula- rensis isolated from most hosts near DPG usually had maximal virulence, but occasion- al isolations had little virulence; (2) carni- vores were readily infected but recovered and did not demonstrate a carrier state; (3) cottontails were su.sceptible to infection; and (4) jackrabbits were more susceptible than cottontails. Klock, Olsen, and Fukushima (1972) obtained eight isolations of F. tula- rensis from Chrysops discalis, Dermacentor panimapertus, H. leporispalustris, and jack- rabbits from four areas near DPG (Delta,^ losepa, Callao, and Gold Hill). Tests of viru- lence showed all eight to be typical Type A (nearctica), which is virulent for laboratory mice and rabbits. EcoDynamics personnel ([Olsen and Dolana] Desert Test Center 1971 Annual Report [USA-DTC 1972]) were mys- tified by the large numbers of seropositive cottontails near Delta because tularemia is imiformly fatal to cottontails. They hypoth- esized that (1) Delta cottontails were some- what resistant to F tularensis as a result of selection pressure, or (2) the detection of the antibody was a nonspecific reaction, or (3) an aviRilent strain of F. tularensis (present in the area) was responsible for the antibody ti- ters. They found support evidence that there probably were specific and protective anti- bodies for cottontail rabbits in the Delta area. Selected details of these supportive data are given later imder "Hosts." On the basis of their findings, the EcoDynamics staff (1972) questioned the validity of the generally ac- cepted concept (based primarily on viru- lence) of types A and B for F. tularensis. Ol- sen and Dolana conjectured further that neither type is limited to specific habitat types nor to specific animal groups. They stated that types A and B allow synthesizing the bewildering array of strains into an epizootiological pattern. No alternative 'Delta, 97 air km (60 air miles) from Dugway, is not physically near, but is environmentally similar with few barriers between the two areas. 110 Great Basin Naturalist Vol. 39, No. 2 method for organization or cataloging was offered. Olsen (1975) reviewed some of the Great Basin surveys by DPG abstracted in the pre- ceding paragraphs. He noted that the major- ity of isolations of F. tuhrensis came from jackrabbits and its primary tick parasite (D. parumapertus), but agglutinins were detected in 22 percent of blood samples from cattle and sheep. Of interest is the decline in rate of recovery of evidence of the tularemia organ- ism in nature from 1954 to 1970. The major- ity of isolations were made earlier with com- paratively little change in field collecting effort. This suggests wide and long-term vari- ation in the amoimt of pathogen circulating in native reservoir hosts and vectors of the area. Epidemiology of tularemia around DPG.- Klock, Olsen, and Fukushima (1973), who described an outbreak of tularemia in 1971 near Delta and Grantsville, Utah, stud- ied the epidemiology of 39 human cases with investigations of vectors, hosts, and incidence of isolations from specimens collected. Nine- teen of these human cases suffered deer fly bites, and another nine reported insect bites by imknown species. The pathogen was iso- lated from hosts and vectors in both areas and adjacent areas such as Skull Valley, Utah. Also, there was an epizootic among la- gomorphs. F. tularensis was isolated from tis- sue of jackrabbits and cottontails. Because of unusually large numbers of midges (Leptoco- nops spp. and Culicoides spp.) observed in 1971, the authors suggested that these might have played a role in the human outbreak. Vectors.— Cox (1965), in a study on F. tularensis and deer flies in the environs of Utah Lake, isolated the organism from Chry- sops spp. in nature for the first time. Experi- mental transmissions by C. discalis was dem- onstrated by Francis and Mayne (1921). Cox's isolation of F. tularensis from C. fulvaster and C. aestuans demonstrated that species other than C. discalis (this common species was only suspect before that time) are vec- tors. Knudsen, Rees, knd Collett (1968) isolat- ed F. tularensis from C. discalis, thus associ- ating the bite of C. discalis with a human case. Other isolations of F. tularensis from deer flies from near the Great Salt Lake, made by the University of Utah Ecology and Epidemiology group, suggested a potential health hazard from Chnjsops spp. Krinsky (1976) pointed out that isolations from deer flies in Utah in 1965, 1968, and 1969 provid- ed evidence of the potential importance of these tabanids in the dissemination of F. tula- rensis, even though they are regarded as short-term mechanical vectors. After these isolations, EcoDynamics researchers made net collections of tabanids and later sugges- ted epidemiologic associations with human cases (Deseret Test Center Annual Reports, USA-DTC 1971-1973). Klock, Olsen, and Fukushima (1973) suggested the same associ- ations (discussed later in this review). Philip (1968) found that another tabanid (Tabanus punctifer, may have been associated with a human case reported near Battle Mountain, Nevada. Parker (1957) and Parker and Johnson (1957) attempted the transmission of F. tula- rensis by fleas in the laboratory. In one at- tempt, three species of fleas were infected, but none of them transmitted F. tularensis to a susceptible host. In another attempt, Or- chapeas leucopus transmitted F. tularensis to a specimen of Peromijscus treui. Extensive studies on transmission of tularemia by ticks, overwintering, and transovarial passage of the pathogen have been conducted at Rocky Mountain Laboratory in Hamilton, Montana (Jellison 1974). Vector studies of tick trans- mission are reported in the University of Utah Ecology and Epidemiology Annual Re- port (USA-DPG 1962b). Otohius lagophilus, removed from a dead jackrabbit, transmitted F. tularensis to cottontails and to a domestic white rabbit (Oryctolagus spp.), the first known transmission by this tick. Twenty-two species of ticks were recorded from DPG and environs, of which five species (Dennacentor andersoni, D. parumpertus, Ixodes kingi, Hae- maphijsalis leporispalustris, and Otohius lago- philus) were infected with F. tularensis (Johnson 1966, Thorpe et al. 1965, and An- nual Reports USA-DPG 1962-1969). It is conceivable that, in the Great Basin, the tick and jackrabbit (D. parumapertus and L. cali- fornicus, respectively) constitute a polyhostal reservoir (Hopla 1974:47). Hosts.— Marchette et al. (1961) demon- strated the susceptibility of locally captured June 1979 Stark: Review of Utah Tularemia 111 wild mammals to tularemia. Eleven species of wild rodents and cottontails were very sus- ceptible. Some species of wild rodents {Ony- chomys leucogaster, Perornyscus maniciilatus, and Neotoma sp.) were very susceptible to virulent strains but were resistant to the avirulent "38" strain. Carnivores {Taxidea taxiis, Vulpes macrotis, and Canis latrans) were readily infected orally; the infection soon was no longer demonstrable, and the an- imals did not become carriers. The authors concluded that tularemia in the Bonneville Basin is a disease primarily of jackrabbits, with the cycle in nature maintained by that host and the vector {Demiacentor paruma- pertus). The authors also concluded that Lepus spp. were more susceptible than Syl- vilagus or Oryctolagus. The published con- clusions of these authors differ from the gen- eral concept throughout the Annual Reports (USA-DPG 1964-1969) that jackrabbits are so susceptible that they play a minor role in maintenance. The snowshoe hare (L. ameri- caniis; European counterpart— L. timidus) is less susceptible than jackrabbits or Minnesota Sylvilagiis spp. to infection with F. tularensis (Green 1943, Green, Larsen, and Bell 1939). Marchette et al. (1961) noted that L. c. deserticola was more susceptible than L. c. te- xianiis. About half of 19 cottontails (S. aiidobonii) captured near Delta showed no demonstrable levels of antibody to F. tularensis (EcoDy- namics Inc. USA-DTC 1972). However, 9 survived the challenge inoculation of 83 cells of F. tularensis; they had developed high lev- els of antibodies. Subsequently, an isolate from cottontails from Delta was tested on other cottontails collected from Delta and Gold Hill. All of the experimental hosts from Gold Hill died, suggesting that some of the Delta cottontails were naturally "immu- nized" with an avirulent strain of F. tula- rensis. This natural avirulent strain may have been responsible for previously observed seropositive cottontails (31 percent of 62 specimens) collected near Delta in 1972 (USA-DTC 1972). EcoDynamics concluded that Delta cottontails may possess some in- nate resistance to certain strains of F. tula- rensis. However, much more work needs to be done to clarify the relation, if any, be- tween lethality of various strains and the re- sponse of partially resistant hosts (such as lab- oratory rats and cottontails). The EcoDynamics group (USA-DTC 1972) reported that 54 carnivores (41 percent) had significant levels of hemagglutinins for F. tularensis. This group of carnivores included 52 bobcats {Lynx rufus), 19 badgers {Taxidea taxus), 8 skunks {Mephitis mephitis), 19 coy- otes {Canis latrans), 30 kit foxes {Vulpes macrotis), and 2 domestic cats {Felis domes- ticus). The positive specimens came from the 13 collecting areas around DPG. A more sen- sitive hemagglutination technique was devel- oped in the early 1970s. Therefore, the high- er percentage of serum samples with agglutinins found from 1970 to 1972 (11 per- cent), may be expected to have higher (though not significantly higher) values when compared to those calculated from tube ag- glutination testing from 1960 to 1969 (roughly 8 percent). The two tests (tube ag- glutination, later replaced by micro- agglutination, and hemagglutination) are dis- cussed later under "Methodology." The phenomenon of carnivorism among rodents is well known in literature for both dissemination and maintenance of plague and tularemia. Vest and Marchette (1958) fed car- casses of 119 rodents {Perornyscus spp.) in- fected with F. tularensis to 11 species of ro- dents (squirrels, heteromyids, and cricetids). In all cases, every rodent that ingested in- fective flesh contracted tularemia. Some spe- cies (mostly heteromyids) were reluctant to feed on flesh, but did so when starved. The extent to which wild rodents supplement their natural diet with flesh was not deter- mined. Cabelli, Ferguson, and McElmury (1964) and Cabelli et al. (1964) reported that the mourning dove {Zenaidura macruora) is rela- tively resistani- to F. tularensis, but the au- thors speculated that fecal transmission could be significant in dissemination of tularemia to susceptible birds. It is of interest that, when a Schu strain of F. tularensis was administered to several species of oceanic birds by several routes, few transmissions from infected to susceptible birds were observed. Francisella tularensis, however, was found in the bird ex- creta although no other evidence of disease (bacteremia) was noted. The brown noddy {Anous stolidus), whitecap noddy (A. min- 12 Great Basin Naturalist Vol. 39, No. 2 utus), white tern (Gygis alba), and sooty tern (Sterna fuscata) were susceptible to infection. Infection in these birds was achieved by res- piratory or cutaneous pathways, rarely by the oral route. Host immune system.— Thorpe and Mar- cus (1962, 1964a,b, 1965a,b,c, 1966, 1967) and Thorpe, Sidwell, and Marcus (1964) re- ported on the implication of phagocytosis in F. tularensis infections. They determined that the phagocytic system of the immune mecha- nism functions as importantly as the antibody system for resisting and overcoming in- fections. Studies of F. tularensis in soils and ON fomites.— At DPG research and testing was conducted on F. tularensis to charac- terize its aerosol stability and persistence in soils and on fomites. Many observations and experiments were conducted by DPG Life Science Division that were apart from studies by contractor groups to DPG such as the University of Utah and EcoDynamics, Inc. and in-house field work by the Environmen- tal and Ecology Branch. Most specific studies on F. tularensis were highly specialized and printed as numbered reports (other than USA-DPG 1951-1969, USA-DTC 1971-1973, USA-DPG 1976, 1978, and University of Utah contract reports). The primary purpose of field-related work (referenced above) was to study the nature and geographic distribu- tion of tularemia occurring naturally in the DPG area. An additional purpose was to monitor the potential for intrusion or estab- lishment of tularemia relative to test activi- ties. Both laboratory and field studies of aero- sols were conducted from the early 1950s through the late 1960s. Many facts were re- ported concerning F. tularensis in nature. Unfortunately, this information was not read- ily available to the scientific community at large. Many of these documents are now available through interlibrary loan. An experiment by Thome (1966) is per- tinent to this presentation. Three pre- dominant soils of the DPG area (sand dune, clay flat, and salt flat) were inoculated with F. tularensis. The pH of the three soil types varied but little (7.5-7.9). In general, sand dunes were the least favorable for preserva- tion of F. tularensis, probably because this soil dried out faster than the other two. High moisture in the soils at the time of in- oculation was the single most important fac- tor for survival. Survival of the organism was better below the surface. Subsequent wetting increased survival. Exposure to sunlight was deleterious. The maximum times after in- oculation that tularemia organisms could be recovered from the soil were 90 days in win- ter and 35 days in summer. When conditions were adverse (as in summer heat), survival was as short as a few hours. An interesting problem has been reported from USSR by Olsuf'yev and Rodnev (1960). Tularemia infection of humans is acquired by the pulmonary route while harvesting cereal crops. Crops were apparently contaminated by infected F. t. palearctica, and human in- fection resulted from inhalation of dust raised into the air from contaminated straw and grain. These conditions are reported to have caused mass infections in people processing the harvest (Spendlove 1974). Popek et al. (1969) reported that washing sugar beets con- taminated by infected rodents in a sugar beet works in South Moravia created aerosols that infected 237 people over four harvesting sea- sons. To date, similar events have not taken place in the Great Basin. Waterborne tula- remia did infect human beings in Oregon during a meadow mouse population explo- sion (Jellison, Bell, and Owen 1959). A poten- tial for airborne infection of tularemia exists throughout the Great Basin whenever thresh- ing or harvesting operations occur. Methodology Serology.— With the basic technique of Alexander, Wright, and Baldwin (1950), the staff of EcoDynamics and their predecessors developed a highly sensitive hemagglutina- tion (HA) test. Dr. Bruce Hudson at the Pub- lic Health Service, Center for Disease Con- trol, Fort Collins, Colorado, assisted in preparing lipopolysaccharide extract of F. tularensis for sensitizing sheep cells. This method is currently used at DPG. Tube ag- glutination (TA) was replaced by micro- agglutination (MA) in 1977. Hemagglutina- tion (HA) was first tried by EcoDynamics in 1970, and this test replaced TA in 1972. Duplicate samples were tested in 1973 and 1978. According to the 1973 Annual Report June 1979 Stark: Review of Utah Tularemia 113 (USA-DPG 1976:89-94 and Table 44), testing by untanned sensitized sheep erythrocytes was more sensitive to the presence of anti- bodies to F. tularensis than was testing by ag- gkitination of a stained antigen (MA) by anti- bodies in unknown samples. The polysaccharide sensitized sheep cells are more likely to yield false positive tests. Today the more sensitive HA test is used as a screen and second confirmation, and the less sensitive MA test is used as a final con- firmatory test. In all replicates MA and TA tests gave similar results and TA has been dis- continued. Recovery of pathogen from infected ARTHROPODS.— Improvements in techniques are presented in the 1973 Annual Report (USA-DPG 1976). Though the passive HA micro-test became firmly established, dis- cussions of other methods in earlier Annual Reports (USA-DTC 1971-1973) merely al- lude to more recently developed methods. One of these is the hemolymph test (Burg- dorfer 1970), in which the distal end of a tick leg is cut and the hemolymph collected on a slide, stained with immunofluorescent or oth- er dye, and examined. This method was dis- cussed in some detail in the 1971 Annual Re- port (Deseret Test Center USA-DTC 1972). However the test has not been used at DPG. Skin test for evidence of tularemia.— Cutaneous allergic reactions have been de- veloped for the diagnosis of tularemia in ro- dents and rabbits (Lundgren, Marchette, and Nicholes 1961, 1962). Cutaneous sensitivity was elicited by inoculating as few as 10 F. tularensis cells; sensitivity was demonstrable for 25 to 53 weeks, depending on species. This diagnostic test aids in establishing pre- vious infection with tularemia for wildlife species. Buchanan, Brooks, and Brachman (1971) reviewed the use of skin tests for hu- man clinical identification of tularemia and epidemiologic studies. Human reactors may remain hypersensitive for as long as 40 years. Antigen prepared for a human skin test (but suitable for use on hosts such as jackrabbits) is available at the Center for Health and Envi- ronmental Studies at Brigham Young Univer- sity. Immunofluorescence.— Karlsson et al. (1970) and Karlsson and Soderlind (1973) re- viewed the use of immunofluorescent (IF) technique in identifying 54 strains, 4 from man and 50 from natural hosts (mostly hares and ticks) in Sweden. They found that once techniques have been established in a given laboratory, histopathologic examination was simplified, and the hazard of laboratory- acquired infection was reduced. The IF method was successful for identifying tula- remia in decomposed material. Cutaneous al- lergic reactions and IF techniques are two promising methods for identifying current tularemia activity in human and other mam- mal populations around DPG. A technique for sampling tabanids for F. tularensis.— Knudsen, Rees, and Collett (1968) described a trap designed to obtain large numbers of deer flies and other tabanids for surveys for the tularemia organism. The trap is pyramidal, about 1 m high and 1 m across the base. About midway up the triangle face on either side is a rectangular flapped opening for flies to enter. Once in- side, the flies can enter a funnel (20 cm diam- eter) against the tip of the pyramid. The top of the funnel is closed but has small openings around the perimeter. Some of the flies inside the funnel may fall through the narrow end, through a plastic tube into a styrofoam box beneath containing 4 to 8 kg of dry ice, which freezes the flies. The sublimation of carbon dioxide also serves as an attractant and is the only bait used. During one season (1968), Knudsen, Rees, and Collett trapped 1,248 deer flies (C. discalis) from marshes bordering the southeastern shore of the Great Salt Lake. Isolates of F. tularensis were ob- tained from flies trapped in this manner. Recent and Current Projects During 1974 and 1975 personnel at DPG further improved the hemagglutination test, which had proved valuable for the staff of EcoDynamics. Lipopolysaccharide antigen was prepared for serologic examination of wildlife and livestock serum specimens col- lected around DPG. The macroagglutination test has been replaced by a micro- agglutination test in accordance with Massey and Mangiafico (1974). Owen (1974) de- scribed morphology and characteristics of F. tularensis and provided information useful in growing and identifying isolates. 114 Great Basin Naturalist Vol. 39, No. 2 With these recently improved techniques it has been reconfirmed that low levels of an- tibody occur in serum samples taken from ro- dents and lagomorphs and a higher incidence occurs in serum samples from carnivores. As presented in preceding annual reports (USA- DPG 1951-1969), sera from a few wild horses at DPG were nearly all positive, but generally antibody response in livestock var- ied. A greater percentage of sheep seem to possess tularemia agglutinins after a stay on a distant summer range than sheep bled during the winter around DPG. Curiously, speci- mens of deer serum contributed by hunters in 1973 possessed no demonstrable antibody, though response in deer had been high during some previous years. Incidence of tularemia in wildlife around DPG will be reviewed in future publications. The first will deal with jackrabbit population changes and tularemia. There has long been an assumption that population reduction of jackrabbits is somehow associated with out- breaks of tularemia in nature. During the last 13 years quantitative data have been collect- ed around DPG which support the general observation that the jackrabbit is subject to large changes of a cyclic nature. While dying jackrabbits infected with F. tularensis have been found, the organism or the disease it produces has not been proved responsible for a widespread decline in jackrabbits. For ex- ample, Philip, Bell, and Larsen (1955) record- ed infected Dermacentor parumapertiis from infected jackrabbits during a peak in the pop- ulation cycle of these lagomorphs in Nevada. Firm evidence that tularemia was responsible for eventual decline of the population was not demonstrable. There appears to be simul- taneous association of pathogen activity (e.g., evidence of antibodies) from cattle, sheep, carnivores, cottontail rabbits, jackrabbits, and rodents. There also seems to be an inverse correlation with jackrabbit density, particu- larly the marked decline of jackrabbits in the DPG area (definitely evident in 1973). At that time there was an apparent increase in tularemia activity. This event followed rather closely the increase of activity of F. tula- rensis in all indicator species during the early 1970s. Epizootiologic and epidemiologic studies were conducted in California (outside the Great Basin) by Lane and Emmons (1977). They concluded that human cases have grad- ually decreased since 1927, and they theorize that the cause was urbanization and selective pressures that have resulted in a decreased virulence of organisms maintained by reser- voir species. A gradual decrease is not evi- dent from data on human tularemia collected from Utah, but urbanization correlates with a decrease of incidence of tularemia as in Cali- fornia. There is a periodic resurgence in time that may be more marked in Utah than in California. DiSTRIRUTION OF ReCENT HuMAN CaSES The Communicable Disease Section of the Utah State Division of Health has kept re- cords of reportable diseases, including tula- remia, for many years. With their permission, recent records representing current condi- tions have been tabulated and are presented as part of Figure 1. Thorpe et al. (1965) and Jellison (1974) have assembled records of tularemia cases through 1964. Jellison's for- mat has been used here with additional de- tailed information included. In Figure 1 the number in each county on the upper right is 1 percent of the human population in 1977. The number on the up- per left is the total cases of tularemia report- ed from 1941 through 1977 (37 years). The larger figure in the center of the county is the number of tularemia cases per 100,000 persons per annum, averaged for 37 years. The ratio between the total number of cases in each county and the 1977 population of each county was analyzed with a posteriori test by simultaneous testing of the homoge- neity of sets of replicates for goodness of fit (G-Statistic, Sokal and Rohlf 1969). Ten sets of samples (under "maximum nonsignificant ranges" in Table 1) were homogeneous; that is, no ratio within each of these sets was sig- nificantly different at the 5 percent level of confidence. The counties which are indicated as significantly higher under "exclusive ranges of homogeneous ranges" overlap with no ratio included in the low group. Ratios in Table 2 are expressed as average cases per 100,000 per annum on the map. Three mral counties, Daggett, Millard, and Rich, had val- ues above 15 per 100,000. Seven more rural June 1979 Stark: Review of Utah Tularemia 115 Large number in center of county - incidence per 100,000 human "^ * " ,^°_^ population. ;?.v.vA\v^v.':\V7Avrv:v7.v.' fiT' ;22 "aal^^j^ Small number in upper left ot fc-82iS^ county = no. cases human tularemia. Small number in upper right of r//.v/.v.v.\\v.v.\v.v.v.v.v.v.v.v.v.-.v.v.v.v.v. ^^^ county = human population x luu. -' ' ' " <^S ITTAFI ^•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.'.•.■.•.•.•.' WEBER i.-! v>!v!v!XvXv;v!->!v!v!\ v!v! v!v;v!-! X-'' 0.53-i.^ <= i,&^ N\y /A V. V. V. V Y." •^- '^ •••••••••••••» 17 1220 1 0 .--•;•. ic !v!v!v!'!v!v!v!v.' o a v i s ■ n W Y 0 M 'ndeterminate N G .y.y .•.•.'.•.•.•.'.*.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.•.'.•.•.•.•.•.'.•.•. . 1 1 6:_5-2 0 os '.'■•.'•:'-''i be ^^^■>^:''^•^•'^^.'^^'^\"^^ !7.03? •.•.•.•••.•.•.•.•.•.•.•.•••. T 0 0 E L E\*--./\ '5^°°>.., ;••:•:•:■ 3. 86; ;.;.;.;.;.;.;.;.%; salt >:•/.;:•.: !v!v!v!v!vi lake ■X-X-X-XvX 0.59,^'- 79 -■ i720' 2.19 ,24 .V/.V.) ^, .•/i .,5 3 ;.v ,/.;.;.;/.- .J u •Xv.e.ir 10.62? e'"!%\"\X\\X\X%XvX' ^ ^ "•' '^ •x':"*2VS?:§xc^ , „■v■E^xv:v^A^^^ "^ ..^ .^.'Mv\^*-x^^^^^^^^^^ ^^^^Mf^^^^^^^PvXi^Vos/':!/^^^ •X\vx-XvXvXvXv •■•■•*•'•'- •'••"■^^"V^^^^^ •XvXv'*'**-''- ^^'^^^ " A ■ " V ■ ■ ■ F " I " " E ■ ' L D ^Xv>A^^^v;^^•X^^^^^^ Fig. 1. Incidence of human tularemia, 1941 through 1977. 116 Great Basin Naturalist Vol. 39, No. 2 counties, Beaver, Carbon, Duchesne, Emery, Sanpete, Uintah, and Wasatch, had values above 7 per 100,000. Counties with the low- est values (less than one per 100,000), Davis, Weber, Morgan, and Salt Lake, are located along the Wasatch Front, which has dense human population (except for Morgan Coun- ty). Washington County also has a very low value. The average for the entire state of Utah (1941 to 1977) is 1.66 cases per 100,000 per annum. Figure 2 shows the actual number of known cases in Utah by year from 1960 through 1977. Two periods of increase above average occurred during the years 1961 through 1964 and 1969 through 1971. Values less than 10 were experienced from 1965 through 1968, and the high of 1971 was fol- lowed by a series of values which were low- est for the longest departure from the aver- age for 1972 through 1977. Distribution of cases by month for the 17 years preceding 1978 is shown in Figure 3. These cases represent date of report, not date of onset or contact with sources of infection. In most cases contact would occur about a month earlier, making the lowest three Table 2. Homogeneity of sets of replicates tested for goodness of fit. Ratio: Maximum Cases Nonsignificant Pop. Ranges Exclusive Ranges of Homogeneous Ratios Daggett .01000 Millard .00634 Rich .00562 Uintah .00393 Beaver .00357 Duchesne .00327 Wasatch .00314 Emory .00312 Carbon .00301 Sanpete .00277 GaFfield .00257 Juab .00226 San Juan .00223 Summit .00143 Box Elder .00116 Kane .00111 Grand .00087 Sevier .00083 Tooele .00081 Piute .00077 Iron .00063 Wa^ne .00059 Utah .00046 Cache .00045 Salt Lake .00022 Washington .00022 Morgan .00021 Weber .00020 Davis .00014 Indeterminate months January through March, and highest three months July through September. Review and Conclusions Tularemia organisms were probably pres- ent in nature and in man around the turn of the century in Utah and perhaps earlier. Francisella tularensis in Utah is ubiquitous in terms of habitat. Type B (F. t. palaearctica) typically occurs as an aquatic infection and may be transmitted from natural sources (in- cluding water and aquatic mammal hosts) to humans and to nonaquatic hosts by ingestion, contact, aerosols, and several arthropod vec- tors (Jellison, 1974). Type A (F. t. tularensis), the more virulent form, which usually occurs in nonaquatic hosts and their arthropod vec- tors, is more widespread; its distribution overlaps the aquatic type. Human cases have occurred in all counties of Utah. The Delta area has sustained repeated human tularemia infections in which the deer fly has served as a major vector. Although F. tularensis is highly invasive, it is fragile and difficult to cultivate and main- tain. Strains of greater and lesser viRilence occur in nature, apparently simultaneously at the same site and even in the same host or vector. Despite well-documented strain dif- ferentiation, only two subspecific designa- tions are accepted currently. Classification of tularemia organisms is a complicated subject. No coordinated effort by researchers has been made to keep all of the strains, and few attempts have been made to verify strains available. In Utah only DPG maintains any identified strains. Although numerous variant strains occur in nature, workers have agreed to accept types A and B (or their counterpart names in refer- ences to this disease). While these concepts are not necessarily germane to cUrrent stud- ies, it is necessary to recognize the problems that numerous strains, which are always pres- ent in natvire, pose in everyday studies. Recent contributions made by personnel at DPG include evidence that the pathogen ex- ists in virtually all areas where specimens are collected regularly, that nonaquatic hosts and their vectors seem to harbor the more viru- lent Type A (F. t. tularensis), and that rodent hosts seem involved .superficially. The role of June 1979 Stark: Review of Utah Tularemia 117 jackrabbits in maintenance and transfer of tularemia is poorly understood. Serum speci- mens with high titers are most frequently found among carnivores, which apparently do not develop overt symptoms but maintain antibodies to F. tiilarensis for a long time, and livestock that are exposed to vectors. As with jackrabbits, the part that carnivores and livestock have in epizootiology of tularemia is unclear. Although some birds are suscep- tible, the pathogen passes from birds with difficulty. They probably play no role in. epizootiology in the Great Basin. Levels of serum agglutinins among various species of mammals correspond well both geographi- cally and in time. As the antibody levels rise slightly ix} rodents and lagomorphs, a con- comitant increase is found in serum samples of carnivores and livestock. Tests by the for- mer Plague Laboratory in San Francisco and at DPG indicate that fleas are probably not vectors. To improve our knowledge of epizootiol- ogy of the disease, one needs a clearer under- standing of the role of jackrabbits in trans- mitting tularemia and whether tremendous fluctuations in population density affect fre- quency of transmission. Also needed is a knowledge of strain differences and whether viRilence changes in a given area over a peri- od of time. Finally, a determination should be made of the regression and subsequent oc- currence of tularemia during apparent inter- epizootic times with regard to geographic sites, ecologic niches, and primary hosts and vectors. Because of the variety of conditions under which the disease occurs, Utah appears to be a prime area for study. 40 30 60 65 70 YEAR Fig. 2. Annual occurrence of tularemia in humans in Utah, 1960-1977. 75 77 118 Great Basin Naturalist Vol. 39, No. 2 Acknowledgments References Cited Appreciation is expressed to the following, who offered advice and consultation during the preparation of the manuscript: J. F. Bell, G. L. Choules, G. T. Crane, T. W. Edwards, T. Fukushima, A. T. Hereim, C. E. Hopla, W. L. Jellison, W. L. Krinsky, P. S. Nicholes, C. F. A. Pinkham, T. J. Quan, H. B. Rees, Jr., M. A. Rothenberg, L. L. Salomon, J. C. Spendlove, V. J. Tipton, and C. M. Wheeler. Dr. Pinkham suggested the study of human tularemia by statistical analysis and used a program which he prepared. Initial stages of this accoimt arose from a research grant pro- posal prepared by Dr. Tipton. Research on which this report is based was supported by Department of Army Project IT0061101A91A (In-House Laboratory Inde- pendent Research). AiKiMBAEV, M. A. 1966. Taxonomy of genus Francisella. Izv. Akad. Nauk Kazakh SSR, s. Biol. 5:42-44. Alexander, M. M., G. G. Wright, and A. C. Baldwin. 1950. Hemagglutination using Franciselhi {Pas- teurella) tularensis polysaccharide. J. Exptl. Med. 91:551-556. Allred, D. M., D. E. Beck, and L. D. White. 1960. Ticks of the genus Ixodes in Utah. BYU Sci. Bull., Biol. Ser. 1(4): 1-42. Allred, D. M., G. N. Stagg, and J. F. Lavender. 1956. Tularemia. Experimental transmission of Pasteu- rella tularensis by the tick, Dennacentor parinna- pertus. ].lni.Dis.99:U3. American Public Health Association. 1964. Diagnos- tic procedures for rickettsial diseases. APHA., New York. Chp. 1, p. 54-55. Bell, J. F. 1965. Ecology of tularemia in North Ameri- ca. J. Jinsen Medicine 2:33-44. Bell, J. F., and S. J. Stewart. 1975. Chronic shedding tularemia nephritis in rodents: possible relation to occurrence of Francisella tularensis in lotic waters. J. Wildl. Dis. 11:421-4,30. J F M A M J J Fig. 3. Monthly occurrence of tularemia in Utah, 1961-1977. June 1979 Stark: Review of Utah Tularemia 119 Buchanan, R. E., and N. E. Gibbons. 1974. Bergey's Manual of Determinative Bacteriology. Williams & Wilkins, Baltimore, Maryland. 1246p. Buchanan, T. M., C. F. Brooks, and P. S. Brachman. 1971. 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Phagocytosis and intracellular fate of Pas- teurella tularensis III. In vivo studies with pas- sively transferred cells and sera. J. Immunol. 94:578-585. 1979 Stark: Review of Utah Tularemia 121 1965c. Passively transferred cells and sera in re- sistance to Pastettrella tularensis infections (ab- stract). Bact. Proc. 65:51. 1966. In vivo phagocytosis of Pasteurelh tula- rensis (abstract). Bact. Proc. 66:67. 1967. Effects of streptomycin on intracellular Pasteurella tularensis during in vitro phagocytic studies (abstract). Bact. Proc. (67: ). Thorpe, B. D., and R. W. Sidwell. 1962. Pathogenic organisms isolated from wild animals in western Utah. Proc. Int. N. W. Conf. Dis. in Nature Comm. to Man 17:110-120. Thorpe, B. D., R. W. Sidwell, and S. Marcus. 1964. Cellular aspects of resistance in Pasteurella in- fections (abstract). Bact. Proc. 674:84. Thorpe, B. D., R. W. Sidwell, D. E. Johnson, K. L. Smart, and D. D. Parker. 1965. Tularemia in the wildlife and livestock of the Great Salt Lake Desert region, 1951 through 1964. Amer. J. Trop. Med. Hyg. 14:622-637. U.S. Army Chemical Corps. 1958. Bibliography of tula- remia. U.S. Army Chemical Corps, Fort Detrick, Maryland. 4th ed. 114p. U.S. Army Deseret Test Center (USA-DTC). 1971-1973. Ecology studies in western Utah. Pre- pared by EcoDynamics, Inc., Fort Douglas, Salt Lake City, UT. Contract DAAD-09-70-C-0051. U.S. Army Dugway Proving Ground (USA-DPG). 1951-1969. A study of the ecology and epizool- ogy of the native fauna of the Great Salt Lake Desert. Annual summary progress report. Pre- pared by Utah Univ. Contracts DA-42-007-AMC- 35(R), DA-42-007-AMC-227(R), DA-18-064-CML- 2639, DA-42-007-403-CML-427, DAAD-09-69-C- 0030(R). 1954. Ecology of disease transmission in native animals. Prepared by Utah Univ. Contracts DA- 18-108-CML-4776, DA-18-064-CML-2455, DA- 18-064-CML-3639(R). Semi-annual report. 1 June to 30 November 1954. 1955. Ecology of tularemia transmission in native animals. Annual report 1954-1955. Contracts DA-18-064-CML-2639. Project Nos. 4-98-05-027. 1961. Studies on the ecology and epizoology of the native fauna of the Great Salt Lake Desert: certain immunological responses of wild rodents and laboratory animals following challenge with Pasturella tularensis. Prepared by Inst. Env. Biol. Rsch., Utah Univ. Contract DA-42-007-403- CML-427. Ecology & Epizoology Series No. 60, 66p. 1962a. A review of tularemia. By L. P. Gebhardt, and B. Thorpe. Prepared by Utah Univ. Contract DA-42-007-403-C M L-427(R). 1962b. Summary status report on Pasturella tula- rensis. Prepared by Utah Univ., Special Report No. 88u (R). 1965. Special tularemia report. By D. D. Bode. Prepared by Utah Univ., Ecology and Epizoology Series No. 93. 1966a. The persistence of Pasturella tularensis in soils of Dugway Proving Ground. By D. S. Thorne. DA Proj. No. 1B650212D624-04 USA TECOM Proj. No. 5-7-9043-02. 1966b. Protocol of operational procedures and epizoological research. Prepared by Utah Univ. Contract DA-42-007-AMC-227(R). Utah Univ. Series No. 134:42p. November 1966. 1976. Ecology studies in the Bonneville Basin of west central Utah. Annual progress report for 1973. DPG-FR-xlOOP. 1978. Environmental and ecology branch prog- ress report for 1974 through 1976. Triennial Re- port. DPG-TR-E-157A. Vest, E. D., D. L. Lundgren, D. D. Parker, D. E. Johnson, E. L. Morse, J. B. Bushman, R. W. Sidwell, and B. D. Thorpe. 1965. Results of a five-year survey for certain enzootic diseases in the fauna of western Utah. Amer. J. Trop. Med. Hyg. 14:124-135. Vest, E. D., and N. J. Marchette. 1958. Transmission of Pasteurella tularensis tularensis among desert rodents through infective carcasses. Science 128:363-364. Woodbury, A. M., ed. 1956. Ecological check list. The Great Salt Lake Desert Series, Utah Univ., mim- eographed report 125p. 1964. Pathogen dissemination among biotic com- munities. Div. Biol. Sci. Misc. papers. Utah Univ. No. 4; also Bull. Utah Univ. 55(22): 1-282 (Tula- remia p. 209-218). Woodbury, A. M., and D. D. Parker. 1954. Studies of tularemia, Pasteurella tularensis. Ecology of the Great Salt Lake Desert. Ecol. Res., Utah Univ. Spec. Rpt. No. 2, 14p. CHEMICAL COMPOSITION OF SOME IMPORTANT PLANTS OF SOUTHEASTERN UTAH SUMMER RANGES RELATED TO MULE DEER REPRODUCTION' Jordan C. Pederson- and K. T. Harper' Abstract.— Chemical composition of some major forage plants of mountain summer ranges of southeastern Utah is reported. Grasses are shown to contain significantly less nitrogen, phosphorus, potassium, calcium, and magnesium than either forbs or shrubs. Forbs and shrubs are demonstrated to differ significantly only in potassium content; forbs tested contained more potassium than shrubs. The chemical composition of the forage plants is discussed in relation to mule deer reproductive rates. It is concluded that protein and mineral content of the forage of the two ranges considered (the LaSal and Henry mountains) is less likely to affect reproductive rates than is the relative digestibility of grasses, forbs, and shrubs. The quality of summer forage has been demonstrated to have an effect on body con- dition, general health, and reproductive ca- pacity of deer (Longhurst et al. 1952, Swank 1956, 1958, Julander et al. 1961, Verme 1962, 1963, Yoakum 1965, Nordan et al. 1968, Sni- der and Asplund 1974). If the comparative performance of deer on different ranges is to be understood, the nutritional composition of forages consumed on both summer and win- ter ranges must be known. It is the objective of this paper to provide information on the chemical composition of some important for- age plants of the LaSal and Henry mountains of southeastern Utah. Previous work has shown that mule deer (Odocoiletis hemionus) herds on those mountain ranges differ markedly in respect to reproductive rate (Pe- derson and Harper 1978). Pederson and Har- per (1978) suggested that differences in qual- ity of forage on summer ranges of the two mountain ranges might be responsible for the observed difference in fawn production. Dietary requirements for whitetailed deer {Odocoileus virginianus) have been reported by French et al. (1956), McEwen et al. (1957), Murphy and Coates (1966), Verme (1963, 1965, 1967, 1969), Ullrey et al. (1967, 1971), and Thompson et al. (1973). Although there is little habitat overlap between white- tails and mule deer, it seems likely that nutri- tional data compiled for whitetails will have some relevance for mule deer nutrition and management. Studies on mule deer food habits and nutrition have been made by Ha- gen (1939), Bissell et al. (1955), Swank (1956), Umess et al. (1971), McCullock and Umess (1973), Robinette et al. (1973), Urness et al. (1975), and Pack (1976). Recent investigations have demonstrated that Rocky Mountain mule deer herds resi- dent on the LaSal Mountains produce about 40 percent more fawns per 100 does than do Henry Mountain herds (Pederson and Harper 1978). In an attempt to identify factors re- sponsible for observed reproductive differ- ences, range condition, diseases, and para- sites, as well as late winter body conditions of the deer, were evaluated over an eight-year period on the two mountains. Results demon- strated that deer averaged larger per sex and age class on the LaSals than on the Henry Mountains, but neither diseases nor parasites differed significantly between herds. Like- wise, deer taken from the two herds could not be shown to differ significantly in respect to body condition in late winter. Winter ranges used by the two herds were similar in respect to both composition and production. In contrast, annual summer forage produc- 'Support provided by the Utah Division of WildUfe Resources and the Utah Division of Water Resources. 'Utah Division of Wildhfe Resources. Springville, Utah 84663. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 122 June 1979 Pederson, Harper: Utah Ranc;e Plants 123 tion on the LaSals averaged almost 65 per- cent greater than on the Henries (Pederson and Harper 1978). Composition of the forage crop on summer ranges also differed sharply between mountains, with LaSal ranges being dominated by good-quality forbs while Henry Mountain summer ranges were heavily domi- nated by shnibs and grasses (Pederson and Harper 1978). In this report, the hypothesis that forage conditions on the summer range are respon- sible for the greater productivity of the LaSal Mountain deer herd will be investigated. Em- phasis is concentrated on nutritional charac- teristics of major forage species of summer ranges utilized by the two herds. We greatly acknowledge the help and sup- port given by the following Utah Wildlife Resources personnel: H. D. Stapley, N. V. Hancock, K. L. Nelson, A. Fleck, L. Hall, and N. K. Larsen. Thanks is extended to A. C. Rancher for his help with the statistical anal- ysis of our data and to G. Cronin for her crit- ical review of this manuscript. Study Areas The stvidy areas are in southeastern Utah. The LaSal Mountains are east of Moab in Grand and San Juan counties. The Henry Mountains lie southwest of Hanksville in Wayne and Garfield counties. The two areas are about 117 km apart. Both are laccolithic mountains of similar geologic age (Butler 1920, Hunt et al. 1953). Precipitation aver- ages somewhat higher (about 10 percent) on the LaSals than the Henries for comparable vegetation zones (Pederson 1970). The LaSal herd unit encompasses approx- imately 221,374 ha. The highest point on the LaSals is Mount Peale at 3,876 m elevation. The Henry Mountain area includes approx- imately 72,886 ha; the highest point on the range is Mount Ellen at 3,500 m. site, an area about one hectare in size was se- lected in aspen forest. A composite sample of the surface 1.5 dm of soil and the five most common grass, forb, and shnib forage species were collected for subsequent analysis. All aboveground parts of grasses and forbs and current growth (leaves and twigs) of shrub species were taken. Only grasses and forbs in flowering condition were collected; most shRib species were not in flower. Individual plants of each species were collected until a composite sample of over 75 g fresh weight was acquired. Samples were lightly packed in paper bags and oven dried at 80 C within 48 hours of harvest. Soil samples were air dried, passed through a 2 mm sieve, and delivered to a commercial, analytical laboratory. Soil pH was deter- mined on a 1:1 soil-to-water mixture with a glass electrode meter. Texture was deter- mined using the hydrometer method, and ex- changeable cations were extracted with neu- tral ammonium acetate and determined by atomic absorption procedures. Nitrogen was measured using micro-Kjeldahl apparatus. Soil organic matter was determined by loss- on-ignition. Dried plant samples were ground through a 40-mesh sieve in a standard mill. Samples were stored in glass containers until the anal- yses were completed. Estimates of crude pro- tein were based on total nitrogen as deter- mined by micro-Kjeldahl procedures. All other elements were determined from ash, us- ing standard atomic absorption and color- imetric techniques (American Society of Agronomy 1965). Differences in chemical composition be- tween species belonging to different life form groups (i.e., grasses, forbs, and shrubs) on the same mountain range and between life forms on different mountain ranges were deter- mined using analysis of variance and Duncan multiple range test. Statistical procedures fol- low Snedecor and Cochran (1967). Methods and Procedures During late July 1976, a representative ' range site in the aspen zone of each moun- tain site was visited. The study site on the LaSals was at Warner Lake; the Henry Mountain site was at Nasty Flats. At each Results Soils of the two study sites are both of loamy texture (Table 1). Soils at the LaSal study area were slightly more acidic than those at the Henry site. Soil organic matter and nitrogen content were somewhat higher 124 Great Basin Naturalist Vol. 39, No. 2 at the LaSal Mountain study area; both varia- bles probably reflect a somewhat better mois- ture balance at the LaSal site. Soils from the Henry Mountain site have considerably high- er phosphorus content and generally higher exchangeable cation levels than the LaSal soils (Table 1). Chemical composition of the current year, aboveground growth of major species, and current-year twig growth of important browse species of the summer ranges of the LaSal and Henry mountains are reported in Table 2. Average values for grasses differ sig- nificantly between mountain ranges. Species from the LaSals contain more nitrogen, po- tassium, calcium, and magnesium. Phos- phorous content of grasses did not differ sig- nificantly between mountain ranges. Shrubs of the two mountain ranges differed signifi- cantly for nitrogen only, with LaSal shrubs averaging 40 percent more nitrogen than the Henries. Elemental content of plants was strongly correlated with soil content of the same element for nitrogen and phosphorous only. Other elements showed little correla- tion between amounts in plants and associ- ated soils. The elemental content of grasses (all spe- cies polled) was significantly lower than for either forbs or shrubs for all elements tested. Few significant differences in chemical com- position could be demonstrated between forbs and shrubs. Only the potassium content of forbs could be shown to differ significantly from that of shrubs— potassium averaged 138 percent higher in forbs (Table 2). The average contribution of grasses, forbs, and shrubs to the forage crop of summer ranges of the LaSals and Henries is shown in Table 3. Furthermore, several analyses of the relative preference of mule deer for grasses, forbs, and shrubs during the summer season have been reported (Table 4). Because esti- mates of chemical (Table 2) and botanical (Table 3) composition of the summer range forage crop of the LaSal and Henry moun- tains are available, it is possible to combine those data with feeding preference informa- tion for mule deer (Table 4) and obtain esti- mates of the chemical composition of the summer diet of deer on the two ranges. Assuming deer select grasses, forbs, and shrubs in the proportions reported by any au- thor in Table 4, regardless of the abundance in the vegetation of plants in each life form category, the composition of the diet can be estimated for any element in either of the study areas. For example, to estimate the amount of phosphorus in diets of LaSal mule deer, assuming a feeding preference such as that reported by Smith (1952) in Table 4, sum the products of (1) average percent phosphorus in LaSal grasses times the pro- portion of grasses in mule deer diets reported by Smith, (2) average percent phorphorus in LaSal forbs times the proportion of forbs in the diet, and (3) the average percent phos- phoRis in LaSal shrubs times the proportion of shnibs in the diet. For this example, we es- timate that mule deer diets on the LaSals should contain .18 percent phosphorus. The chemical composition of four alterna- tive diets has been estimated in Table 5. Three diets are based on feeding preferences reported by Smith (1952), Morris and Schwartz (1957), and Trout and Thiessen (1968). The fourth diet is based on the as- sumption that the deer select grasses, forbs, Table 1. Chemical and physical characteristics of soils on the LaSal and Henry mountains study areas. Mountain Sand Silt range PH % % LaSal Moun- tains 6.3 48 46 Warner Lake Henry Moun- tains 6.7 50 38 Nasty Flats Analysis Organic Clay matter Nitrogen Phosphorus Potassium Calcium Magnesium ^ % % ppm ppm ppm ppm 14.0 417.( 42.4 420.0 2090.4 105.1 fune 1979 Pederson, Harper: Utah Range Plants 125 and shrubs in exact proportion to their abun- dance in the vegetation. The results show chemical composition of diets differs relative- ly little regardless of the assumption used (Table 5). Furthermore, estimated chemical composition of diets does not differ radically between mountain ranges. Even the lowest estimates for each dietary constituent studied appear to be within safe limits for good ani- mal health (Dasmann 1971, Morrison 1961). The calcium /phosphorous ratio also seems to be within normal limits (Table 5). Discussion The results indicate that differences in plant chemistry between the two mountain ranges for the elements considered in this pa- per are probably not responsible for the ob- served reproductive differences between mule deer herds resident on the two moun- tain ranges. However, it cannot be concluded that summer range forage is not responsible for the observed differences in deer repro- duction. The digestible energy content is un- known for the species considered in this re- Table 2. Chemical composition of current year, aboveground growth of some major forage species of the summer ranges of the LaSal (L) and Henry (H) mountains of southeastern Utah. Specimens for analysis were collected on 26 and 27 July 1976. Composition values have been averaged by plant life form group and mountain range. Species Nitrogen Phosphorus Potassium Calcium I Magnesium L H L H L Percent H L H L H Grasses Agrapyron trachycaulum 1.51 0.91 0.12 0.12 1.88 0.89 0.26 0.27 0.07 0.08 Bromus cannatus 1.79 — 0.17 — 2.60 — 0.44 — 0.12 — Carex geyeri 1.16 _ 0.12 _ 1.81 _ 0.34 _ 0.13 _ Elymus glaucus 1.82 _ 0.13 _ 2.61 _ 0.35 _ 0.09 _ Festuca ovina - 1.30 - 0.11 - 1.06 - 0.26 - 0.07 Festuca thurberi - 0.99 - 0.10 _ 1.75 - 0.22 _ 0.07 Sitanion hystrix - 1.15 _ 0.11 - 1.64 _ 0.17 - 0.06 Stipa Columbiana 1.19 1.11 0.12 0.13 1.82 1.55 0.33 0.33 0.08 0.08 Average by mountain range 1.50* i.ogf' 0.13a O.IP 2.14a 1.38^ 0.35a 0.2^ 0.10* 0.07^ Pooled average 1.305^^ 0.123' 1.762'' 0.300P 0.085^^ FORBS Chenopodium fremontii - 3.14 - 0.25 - 7.90 - 1.18 - 0.76 Erigeron speciosus 1.77 - 0.18 - 3.35 - 1.01 - 0.15 - Helianthella quinquenervis 1.62 - 0.23 - 4.53 - 1.65 - 0.28 - Hymenoxys richardsonii — 1.54 — 0.20 — 3.05 — 1.29 _ 0.26 Ligusticum porteri 1.55 - 0.19 - 4.74 - 1.34 - 0.25 - Lupinus parviflonts 2.49 - 0.15 - 2.29 - 1.44 - 0.34 - Oxytropis sericeus - 1.92 - 0.18 _ 1.91 _ 0.98 - 0.16 Penstemon watsonii _ 1.12 _ 0.15 _ 1.88 - 1.16 - 0.26 Senecio ambrosioides _ 1.62 _ 0.19 _ 3.24 _ 1.01 _ 0.23 Thalictrum fendleri 1.44 - 0.19 - 2.19 - 0.87 - 0.12 - Average by mountain range 1.77= 1.87? 0.18^ 0.19* 3.42a 3.59a 1.26^ 1.12a 0.23a 0.33P Pooled average 1.820^ 0.192^ 3.509^ 1.195<1 0.281^ Shrubs Artemisia tridentata vaseyana - 1.92 - 0.20 - 2.09 _ 0.49 - 0.15 Populus tremuloides 1.96 1.51 0.13 0.15 1.06 1.38 1.20 0.60 0.20 0.17 Prunus virginiana 2.57 _ 0.30 _ 1.55 _ 1.71 _ 0.33 - Ribes cereum _ 1.19 _ 0.20 _ 1.02 _ 0.94 _ 0.26 Rosa woodsii 1.84 1.86 0.19 0.22 1.09 1.27 1.03 1.59 0.33 0.38 Salix lasiandra 2.59 _ 0.16 _ 1.29 _ 1.33 _ 0.26 _ Symphoricarpos oreophilus 1.86 1.23 0.17 0.18 1.90 2.08 1.10 0.63 0.29 0.23 Average by mountain range 2.16f 1.54b 0.19* 0.19* 1.38* 1.57a 1.27a 0.85a 0.28a 0.24a Pooled average 1.853^ 1 0.191'! 1.474' as long as pronotum; sides almost straight and parallel on basal two thirds, rather broadh rounded behind; surface smooth, brightl) shining, punctures minute, almost obsolete perhaps in obscure strial rows. Declivit\ steep, convex; striae obscurely evident, punc tures more distinct than on disc; interstriae ]j weakly, distinctly elevated, 2 strongly, rathei 1979 Wood: American Bahk Bi:i:tlks 139 narrowly elevated from just below base to just below middle, its narrowly convex crest uniformly elevated, with a row of small punctures, 3 neither elevated nor impressed, with a row of punctures on 1-3 possibly very feeblv granulate. Vestiture restricted to de- clivity, of interstrial rows of subspatulate bristles on all interstriae; about 5-6 on each interstriae, each bristle about one and one- half times as long as distance between rows. Type Locality.— One km southwest of Rincon de Osa, Puntarenas, Costa Rica. Type Material.— The male holotype was taken on 12-VIII-1968, from a Cecropia leaf petiole, by H. Hespenheide. The holotype is in my collection. Dendrocranulus auctus, n. sp. This species is distinguished from limitaris Wood by numerous characters cited below. Female.— Length L8 mm (paratypes 2.3-2.7 mm), 2.7 times as long as wide; color very dark reddish brown. Frons about as in limitaris except tuft of hair less dense. Pronotum as in limitaris except posterior areas reticulate, punctures and their accom- panying granules much smaller, extending to median area. Elytra as in limitaris except strial and in- terstrial punctures rather minute, distinctly impressed and declivity rather strongly im- pressed. Declivity almost smooth, shining, striae obsolete; interstriae 1 slightly elevated, with a row of very fine granules, 2 somewhat strongly impressed, particularly on lower half, with a row of small to feeble granules, 3 higher than 1 and ascending to broadly rounded 4, 3 and 4 each with a row of feeble, minute granules as on 2. Vestiture as in liini- taris except only two-thirds as long on decliv- ity. Male.— Similar to female except frons lesser convex, almost flat on lower half, with rounded tubercles in lateral and dorsal areas, vestiture sparse, declivity much more broad- ly, strongly impressed (shallowly subcon- cave), its lateral and apical margins more abniptly rounded, punctures on declivital striae small, distinct, granules slightly larger. Type Locality.— Rancho Grande, Pittier National Park, Aragua, Venezuela. Type Material.— The female holotype, male allotype, and 48 paratypes were taken at the type locality on 9-IV-197(), 1100 m. No. 407, by me, from the same cucurbi- taceous vines that contained limitaris. The holotype, allotype, and paratypes are in my collection. Dendrocranulus limheUus, n. sp. The four species in this genus named here (excluding modus) are somewhat related to limhatus Blandford and fulgidus Wood, but all are distinguished by the longer, more slen- der declivital setae and by the very different arrangement of long frontal setae on the fe- male. The four are allied to one another, but not to other South American members of the genus presently known to me. Female.— Length 2.4 mm (paratypes 2.0-2.5 mm), 2.9 times as long as wide; color very dark brown. Frons broadly, evenly convex from epis- toma to vertex; surface smooth, shining ex- cept obscure reticulation in some lower areas, punctures rather coarse, deep, close on upper half, finer and subgranulate on lower half of area below upper level of eyes; vesti- ture on lower half of area below eyes of fine, long abundant hair, longest equal to one- third distance between eyes. Pronotum L2 times as long as wide; essen- tially as in limhatus except posterior areas smooth, shining, with no reticulation, punc- tures distinctly larger, rounded tubercles on lateral margins of most punctures slightly larger. Elytra L7 times as long as wide, L6 times as long as pronotum; outline as in limhatus except more narrowly rounded behind; disc as in limhatus except striae 1 slightly im- pressed, strial punctures spaced by 1-2 diam- eters of a puncture, interstrial punctures more widely spaced, declivity narrower, more distinctly impressed on interstriae 2; declivital strial punctures smaller than on disc, interstriae 1 wide, a row of fine, feebly granulate punctures on side next to suture, 2 almost impunctate, 3 with about three fine, rounded granules. Vestiture of interstrial hair, each seta as long as distance between rows, slightly longer toward base of declivity, usu- ally absent on declivital interstriae 2. 140 Great Basin Naturalist Vol. 39, No. 2 Male.— Similar to female except frons al- most flat on lower half, with punctures finer, vestiture sparse, declivity much more strong- ly, more broadly impressed (to middle of in- terstriae 3), its lateral and apical margins more abruptly rounded. Type Locality.— Merida, Merida, Vene- zuela. Type Material.— The female holotype, male allotype, and 25 paratypes were taken at the type locality on 22-IX-1969, 5300 m. No. 1, from Cucurbita, by me. The holotype, allotype, and paratypes are in my collection. Dendrocranulus limitaris, n. sp. This species is distinguished from the very closely allied limbellus Wood by the more extensive female frontal vestiture, by the fin- er punctures on the head, and by differences on the pronotum and elytral declivity cited below. Female.- Length 2.1 mm (2.0-2.3 mm), 2.7 times as long as wide; color very dark brown. Frons as in limbellus except area above pubescence mostly reticulate, punctures much finer, dense brush of pubescence ex- tending from epistoma to upper level of eyes, longest setae equal to almost half distance be- tween eyes. Pronotum as in limbellus except punctures on disc smaller, their accompanying tubercles larger, tuberculate area extending to median area. Elytra as in li7nbellus except declivital in- terstriae 2 ascending laterally, with a row of regularly spaced punctures, granules on in- terstriae 3 absent, vestiture on declivital in- terstriae 2 usually present. Male.— Similar to female except frons less strongly convex (more strongly than male limbellus), with vestiture sparse, declivity more distinctly impressed (about as in female limbellus), with interstriae 2 largely impunc- tate, lateral and apical margins broadly rounded. Type Locality.— Rancho Grande, Pittier National Park, Aragua, Venezuela. Type Material.— The female holotype, male allotype, and 137 paratypes were taken at the type locality on 9-IV-1970, 1100 m, Nos. 407 (type), 421, 422, 423, from cucurbit vines, by me. The holotype, allotype, and paratypes are in my collection. Dendrocranulus modus, n. sp. This species is distinguished from carbo- narius (Ferrari) and guatemalensis (Hopkins) by characters indicated below. Female.— Length 1.8 mm (paratypes 1.6-2.0 mm), 2.5 times as long as wide; color very dark brown. Frons as in guatemalensis except weakly, transversely impressed just above epistoma. Pronotum as in guatemalensis except disc less irregular with punctures smaller than in guatemalensis but larger than in carbonarius. Elytra as in guatemalensis except declivity less strongly impressed (but more so than in carbonarius), declivital interstrial setae much longer than in carbonarius, slightly longer than guatemalensis, each distinctly longer than distance between rows; interstrial punc- tures on declivity very feebly granulate. Male.— Similar to female except frons in- termediate between males of guatemalensis and carbonarius, median third on lower half slightly impressed (not impressed in carbo- narius; in guatemalensis impression extends to lateral margins and higher on frons); de- clivital impression slightly stronger and more extensive than in female. Type Locality.— Merida, Merida, Vene- zuela. Type Material.— The female holotype, male allotype, and 12 paratypes were taken at the type locality on ll-iX-1969, 170 m. No. 1, from Cucurbita, by me. Four para- types are from the same locality, and host, taken 22-IX-1969, No. 18. The holotype, allotype, and paratypes are in my collection. Dendrocranulus pinguis, n. sp. The relationship of this species to acutus, limbellus, and limitaris is more remote. The female frons lacks a dense tuft of hair and the declivity is less distinctly impressed. Female.— Length 2.4 mm (paratypes 2.2-2.4 mm), 2.5 times as long as wide; color very dark brown. |une 1979 Wood: American Bark Beetles 141 Frons broadly convex, a weak, transverse impression between eyes; surface reticulate, rather finely, uniformly punctured, lower margins of most punctures shining, perhaps feebly granulate; vestiture sparse, inconspic- uous; fine, rather short. Pronotmn 1.2 times as long as wide; resem- bling aiictus Wood; disc finely reticulate, punctures very small, distinct, their accom- panying tubercles transverse, with longitudi- nal axis equal to diameter of puncture, trans- verse axis two or three times as great. Elytra 1.5 times as long as wide, 1.4 times as long as pronotum; sides straight and paral- lel on basal two-thirds, slightly tapered then very broadly rounded behind; striae not im- pressed, pmictures moderately coarse, deep, close; interstriae twice as wide as striae, al- most smooth, shining, punctures two-thirds as large as those of striae, rather close. Declivity very steep, broadly convex, feebly sulcate on median half; strial punctures slightly smaller than on disc; interstriae 1 slightly elevated, 2 feebly impressed, 3 as high as 1, interstrial punctures almost as large as those of striae, their dorsolateral margins armed by a granule almost equal in height and diameter to punc- ture. Vestiture abraded on type; on paratypes consisting of erect interstrial bristles, each slightly shorter than distance between rows. Male.— Similar to female except frontal impression slightly more extensive, lateral granules larger, declivity more broadly, more strongly impressed (but still rather shallow). Type Locality.— Bumbum Forest Station, Barinas, Venezuela. Type Material.— The female holotype, male allotype, and six paratypes were taken at the type locality on 29-1-1970, 150 m. No. 276, from a cucurbit vine, by me. The holotype, allotype, and paratypes are in my collection. Hylocurus clarki, n. sp. This species is distinguished from aberrans Wood by the more coarsely tuberculate pro- notal disc, by the smaller discal strial punc- tures, and by the very different male elytral declivity. Male.— Length 2.3 mm (male paratype 2.5 mm), 2.4 times as long as wide; color dark reddish brown. Head about as in aberrans. Pronotum as in aberrans except disc much more strongly reticulate, rounded tubercles conspicuously larger. Elytra similar to aberrans except strial punctures smaller, interstriae slightly wider than striae and marked by more numerous transverse lines, declivity with spines in cir- cumdeclivital ring blunt, conspicuously more strongly projecting, particularly on upper half, degree of projection about equal to width of spine, punctures of declivital face confused, vestiture on circumdeclivital ring conspicuously longer, more slender, setae on declivital face short, of stout hair. Type Locality.— Between Sicabe and San Miguel Ixtahuacan, San Marcos, Guatemala. Type Material.— The male holotype and one male paratype were taken at the type lo- cality on 24-11-1972, from Pinus tenuifolia, by E. W. Clark. The holotype and paratype are in my col- lection. Hylocurus longipennis, n. sp. This species is distinguished by the slender body form and by other characters cited be- low. Though it is more closely allied to hir- tellus (LeConte) than to other known species, the relationship is not close. Male.— Length 2.5 mm (females 2.8-3.0 mm), 3.1 times as long as wide; color very dark brown. Frons with a strong, transverse carina on more than median half midway between lev- el of antennal insertion and upper margin of eyes; surface concealed by pronotum above carina, smooth, shining, with small punctures at sides and below. Antenna about as in hir- tellus except club slightly larger, wider, with sutures slightly more strongly procurved. Pronotum 1.2 times as long as wide; about as in hirtellus except disc longer, more strongly reticulate, with subcrenulate tu- bercles almost twice as large. Elytra 2.0 times as long as wide, 1.7 times as long as pronotum; sides straight and paral- lel on basal four-fifths, rather abruptly, ser- rately tapered to strong, apical mucro; striae not impressed, punctures rather coarse, deep, spaced by diameter of a puncture; interstriae slightly narrower than striae, smooth, shining. 142 Great Basin Naturalist Vol. 39, No. 2 punctures small, close, their anterior margins slightly elevated, more strongly so near de- clivity. Declivity very steep, convex; about as in hirtellus except smoother, more brightly shining, tubercles at base slightly larger, broader, interstriae 3 without tubercles be- low junction with 7, 9 higher and without tu- bercles on its posterior half; costal margin near apex finely serrate. Vestiture much as in hirtellus except interstrial setae at base of de- clivity longer, coarser, strial setae much shorter. Female.— Similar to male except frons without a carina, an indefinite callus in its place, upper surface with indefinite punc- tures and fine, sparse granules, vestiture rather sparse; tubercles at base of declivity finer. Type Locality.— Five km west of El Sal- to, Durango, Mexico. Type Material.— The male holotype, fe- male allotype, and two female paratypes were taken on 7- VI- 1965, 2500 m. No. 41, from a Quercus branch by me. The holotype, allotype, and paratypes are in my collection. ANNUAL ENERGY BUDGETS FOR THREE COMMON RODENT SPECIES IN THE NORTHERN GREAT BASIN' R. Kent Schreiber^''^ Abstract.— Annual energy budgets were calculated for three species of small mammals (Peromyscus nuiniculatiis, Onychomijs leucogaster, Reithrodontomys megalotis) from the northern Great Basin, Benton County, Washington. Lidividual ingestion rates were based on species activity, microclimate regime, coefficient of digestibility, caloric diet, and the cost for reproduction. For males and females, the estimated energy expenditures were: P. maniculatus, 6080, 5891; O. leucogaster, 5714, 6587; and R. megalotis, 4057, 3791 kcal/yr. By comparison, each species on an individual basis processes more energy annually than the more abundant species in the community, Perognathus parvus, but their total contribution to community energy fiow is apparently minor. Integration of these results with other ecological parameters is necessary to develop new hypotheses on the role of small mammal consumers in cold desert ecosystems. Historically, ecologists have studied and compared ecosystems and their component species in terms of density and biomass. However, this approach does not emphasize the impact of each species on the total sys- tem or its relationship to other trophic levels within the system. The concept of energy flow provides such a common factor for com- paring ecosystems and also for evaluating the relative importance and success of the con- stituent populations. In the northern part of the Great Basin common rodent species include the Great Ba- sin pocket mouse (Perognathus parvus), deer mouse (Peromyscus maniculatus), northern grasshopper mouse (Onijchomys leucogaster) and the western harvest mouse (Reithrodon- tomys megalotis). These small mammal con- sumers are representative of an important pathway for energy transfer in a cold desert ecosystem. The bioenergetics of the pre- dominant species, P. parvus, has been dis- cussed in a previous paper (Schreiber 1978b). This paper reports on the energy budgets of the three, less abundant, species. Energy flow through a rodent population can be determined from daily energy require- ments and ingestion rates of individuals dur- ing each season. In this study I calculated in- gestion rates by considering activity of each species in field-encountered microclimates and their resulting metabolic demands. Ad- justments in the ingestion rates were made for the additional energy cost of reproduction and for the energy savings while residing in a nest. Materials and Methods Study Area The study area is in the Hanford Works Department of Energy (DOE) Reservation 19 km northwest of Richland, Benton County, Washington. The 58-year average annual precipitation for the Reservation is 159 mm. Other climatological and edaphic conditions for the general area have been summarized by Stone, Jenne, and Thorp (1972). Vegeta- tion is mostly typical of the Artemisia triden- tata-Poa association (Daubenmire 1970) with the exception of native grass species. Cheat- grass (Bromus tectorum) was introduced into the area over a half century ago and now has replaced native species as the dominant ground cover. Shrubby species present in- clude big sagebrush (Artemisia tridentata), bitterbrush (Purshia tridentata), and two rab- bitbrush species (Chrysothamnus naiiseosus; C. viscidiflorus). 'This study was supported in part by the U.S.-IBP Desert Biome Program (Grant GB 15886 from the National Science Foundation). 'Department of Biological Sciences, University of Idaho, Moscow, Idaho 83843. 'Present address: U.S. Fish and Wildlife Service, National Power Plant Team, 2929 Plymouth Road, Ann Arbor, Michigan 48105. 143 144 Great Basin Naturalist Vol. 39, No. 2 Trapping Although no attempt was made in this study to delineate absolute population num- bers, monthly sampling with snap-traps de- termined species composition, relative popu- lation sizes, and trends. Traps were spaced approximately 3 m apart, with 50 traps per 150 m line. Rolled oats paste was used for bait and traps were normally set for three consecutive nights in each trapping session. Sex, weight, and reproductive status of all captures were recorded. In addition, rodents were live-trapped in peripheral areas for use in laboratory food trials. Microclimate To establish the microclimate regime of the species, temperatures were recorded con- tinuously at the surface and at a burrow depth of 0.5 m by a seven-day, two-pen thermograph. Data were summarized to coincide with monthly trapping sessions. Mean diurnal surface temperature (Tj) was calculated as the average of even-hour tem- peratures from dawn to dusk. Correspond- ingly, mean nocturnal surface temperatures (TJ were calculated as the average of even- hour temperatures from dusk to dawn. Bvir- row temperature (T^) was calculated as the mean of the daily maximum and minimum subsurface temperature. Digestibility Energy content of ingested materials was determined by combustion in a semimicro ox- ygen bomb calorimeter. The coefficient of di- gestibility (digested proportion of ingested food) was measured directly in the laboratory from food intake and indirectly by the ash- tracer method for free-living animals (Schrei- ber 1979). Energy Expenditure Annual ingestion rates were calculated from the activity and resting time of the spe- cies in field-encountered microclimates (sur- face and burrows) and their resulting caloric demands. The additional energy cost of re- production and the energy savings from in- svilating properties of the nest were in- corporated into the calculations. Ingestion rates were calculated using the general model: I = [(E, + EJ + E^] D-i = (E, + Eg^)D-i (1) where I is ingestion rate (kcal/yr), E^ and E^ are energy costs during rest and during activ- ity. En, is their sum (maintenance), E^ fs energy COStS for growth from weaning to subadult, and D is coefficient of digestibility. Additional growth between the subadult and adult stage was considered by using the average adult weight when calculating maintenance energy (En,). During pregnancy and lactation, fe- males incur additional energy demands due to respiration and growth of the embryos. To account for embryonic respiration, gravid fe- males were included in calculations of female mean weight. Ingestion rates for females were calculated as— If = (En, + p Eg^ + w Eg^ + E^) (D-i) (2) —where the coefficient p is mean brood size at parturition (i.e., mean litter size X aver- age number of litters per year), w is the mean brood size at weaning (i.e., mean brood size (p) minus mortality during nursing period), and Eg^ and Eg^ are energy costs for growth from conception to birth and from birth to weaning, respectively. Intrauterine mortality is unknown but probably small and has been ignored in the calculations. Females with ei- ther embryos or placental scars were record- ed as bearing one litter; females with both embryos and scars or scars of an undeter- mined number were recorded as having two litters. For all species I assumed a conserva- tive survival rate of 80 percent for nursing young (Kaczmarski 1966). Energij Costs at Rest: Resting metabolic rates (RMR) for individual species were taken from the literature. Since animals in a bur- row and occupying a nest have lower energy requirements during rest, I adjusted RMR's for this energy conservation by plotting the nesting metabolic rate (NMR) as a regression line based on 0.81 RMR at 1 C and 0.87 RMR at 12 C (based on data reported for the harvest mouse, Pearson 1960). June 1979 Schreiber: Rodent Energy Budgets 145 Energy Cost of Activity: Metabolic rates increase during periods of activity. Estimates of daily and seasonal variations in amounts of activity were based on field-monitored activi- ty of free-roaming mice tagged with a radio- active nuclide (Schreiber 1973), personal communications from other field in- vestigators, and published data (French et al., 1966). The incremental energy demand (EJ during these activity period was calculated from information cited in Chew and Chew (1970). Energy Cost of Growth: Energy cost for growth during a specific stage of devel- opment (Eg) is the product of the weight gain (W) and tissue caloric value (K) divided by growth efficiency (G) during that period of growth, i.e.. E, = (WK)(G- (3) I used the following caloric values for the tissues (K): for the embryo, 0.98 kcal/g fresh weight, based on the average caloric values of five species of newborn rodents (Gorecki 1965, Myrcha and Walkowa 1968, Soholt 1973), and for the unweaned young, 1.39 kcal/g, assuming an average weaning age of 25 days and the mean caloric value of two species of rodents (Myrcha and Walkowa 1968, Soholt 1973). Caloric values for weaned young of individual species are given in the results. Growth efficiencies were taken from the literature and based on average val- ues, G = 13.8 percent for embryos, 15.0 per- cent for unweaned young, and 5.0 percent for weaned young (Kaczmarski 1966, Migula 1969, Drozdz et al. 1972). Results and Discussion Composition and Abundance Snap traps effectively sample small mam- mal populations (Wiener and Smith 1972), and they are particularly applicable for cen- susing large areas. Under ideal conditions the total number of individuals caught in traps is proportional to population density and re- flects the structure of the population (Han- sson 1967, Petticrew and Sadlier 1970). Al- though trapping percentages are not direct estimates of density, they are nonetheless in- dicative of population trends and therefore provide insight into the influence a particular species has in the transfer of energy into the community. Trap lines were operated one night each in September and November 1969 and March, April, and May 1970 and at least three nights per month from June 1970 to May 1971 (Table 1). A total of 1470 rodents was cap- tured in 14,289 trap-days (one trap set for one day). The average monthly effort was 794 trap-days and the overall trapping suc- cess was 10.3 percent. Field observations in- dicated traps placed in dense cover or con- cealed by shadows had somewhat greater success. On other areas of the Hanford Reser- vation general trapping success has been re- ported as low as 4 percent on fire-disturbed grasslands with stony soils (Hedlund et al. 1975) and as high as 44 percent in shrub- steppe habitats with coarse-textured sands (O'Farrell 1975b). A total of four species of rodents was snap- trapped on the study area: the Great Basin pocket mouse, Perognathus parvus; deer mouse, Perornyscus maniculatus; northern grasshopper mouse, Onychomys leucogaster; and western harvest mouse, Reithrodontomys megalotis. Perognathus parvus composed 84.2% of the total catch, with P. maniculatus, O. leucogaster, and R. megalotis comprising 9.4, 3.4, and 2.9 percent, respectively (Table The low trapping success in the fall re- flects reduced surface activity and the post- breeding mortality of P. parvus, the most abundant species. Summer peaks reflect the termination of reproduction in this species and the increased foraging of weaned young. With the exception of R. megalotis, species composition was comparable to small mam- mal populations inhabiting slightly higher elevations on the reservation, where ground cover consists of more native vegetation (O'Farrell et al. 1975). The greater percent of captures of harvest mice on my study area may reflect this species propensity for habi- tats with a mixture of native and introduced vegetation (Black and Frischknecht 1971). Peromyscus maniculatus was the only species taken throughout the year, although it showed considerable seasonal variation in the number of individuals trapped. Perognathus 146 Great Basin Naturalist Vol. 39, No. 2 parvus was conspicuously absent in the cold- est winter months (December and January) and R. megalotis was not trapped in the fall months of September and October. Ony- chomys leucogaster was captured each month except February, but trapping success for this species and R. megalotis never exceeded 1 percent. Other rodent species, including the sagebrush vole, mountain vole, Townsend's ground squirrel, pocket gopher, and bushy- tailed wood rat occur on parts of the reserva- tion but were absent on my study area. Because of the unpredictability of precipi- tation and extremes in temperature, desert rodent populations can demonstrate consid- erable annual fluctuations. The pocket mouse has specifically adapted to this environment (Schreiber 1978a), and the other species, be- cause of their eurytopic habits, are able to survive at low population levels. Even though total population numbers may exhibit large annual oscillations, the proportional distribution of species probably remains stable over the long term. Energy Budgets and Ingestion Rates Energy flow in the individual is a function of the temperature gradient between body temperature (Tg) and ambient temperature (T^). Heat is lost from the body when TaTb. The rate of metabolism is inversely proportional to the temperature gradient at temperatures below thermoneutrality and di- rectly proportional to temperatvires above it. Small rodents, with a relatively large body surface to body weight ratio, gain heat from the environment and dissipation of this heat load against a thermal gradient would re- quire evaporative cooling and subsequent water loss, a luxury desert rodents cannot af- ford. However, these nocturnal animals rarely encounter ambient temperatures that exceed body temperatures (Table 2), so energy expenditure is mainly from thermoge- nesis and activity. Females experience addi- tional demands during pregnancy and lacta- tion. Seasonal changes in the insulatory Table 1. Monthly trapping results for the Hanford Study Area, Benton County, Washington. - Species P- parvus P. maniculatus O. leucogaster R. megalotis Trap Trap Trap Trap Trap success success success success Month days No. (%) No. (%) No. (%) No. (%) 1969 Sep. 200 9 (4.5) 1 (<1) 0 0 Nov. 400 17 (4.2) 1 (<1) 2 (<1) 1 (<1) 1970 Mar.' 280 50 (17.8) 6 (2.0) 0 1 (<1) Apr. 297 50 (16.8) 13 (4.4) 2 (<1) 0 May 298 92 (30.9) 10 (3.3) 3 (1.0) 1 (<1) June 2216 544 (24.5) 39 (1.7) 19 (<1) 8 (<1) July 550 166 (33.2) 7 (1.4) 3 (<1) 4 (<1) Aug. 450 40 (8.9) 8 (1.8) 4 (<1) 1 (<1) Sep. 1050 48 (4.6) 10 (1.0) 5 (<1) 0 Oct. 1648 34 (2.1) 8 (<1) 6 (<1) 0 Nov. 1300 2 (<1) 11 (<1) 1 (<1) 3 (<1) Dec. 600 0 3 (<1) 1 (<1) 3 (<1) 1971 Jan. 750 0 7 (<1) 3 (<1) 3 (<1) Feb. 900 3 (<1) 6 (<1) 0 9 (1.0) Mar. 750 39 (5.2) 3 (<1) 1 (<1) 4 (<1) Apr. 950 60 (6.3) 4 (<1) 0 3 (<1) May 750 47 (6.3) 2 (<1) 0 2 (<1) Aug. 900 37 (4.1) 0 0 0 Total 14,289 1,238 (8.7) 139 (1.0) 50 (<1) 43 (<1) "Heavy rain and strong winds recorded at trapping s 1979 Schheiber: Roi:)ent Energy Budcjets 147 properties of the pelage influence metabolic- rates, but in small mammals this effect is minimal. Therefore annual energy expendi- tures of individuals are primarily the result of reproduction. Rcithrodontoniijs megalotis: This rodent was the smallest of the four species captured and it had a scattered distribution on the study area. Although never abundant, it is an opportunist, which enables it to exploit a va- riety of microhabitats. Pearson (1960) calculated resting metabol- ism (m/ O2 g-i hr-i) in this small cricetid as— E, = 11.41 - 0.27 Tb (Tb<24.5 C) (4) Adding the increment for activity (2.9 ml Og/g/hr, Chew and Chew, 1970) to E^, E^ = 14.31- 0.27 Ta (5) Harvest mice construct elaborate, well- insulated nests which reduce energy ex- pended for thermoregulation at lower tem- peratures. Thus, En = 9.2 - 0.18 Tb (Tb<24.5 C) (6) Harvest mice may be gregarious during the colder months; if so, their metabolic costs would be effectively reduced. Without nest- ing material, huddling can reduce metabolic rates 27-39 percent (Pearson 1960, Trojan and Wojciechkowska 1968). With a nest, huddling reduces energy expended in heat production by about 13 percent (Grodzinski and Gorecki 1967) and significantly lowers food consumption (Gebczynska and Geb- czynski 1971). I used the latter figure (13 percent) to determine the savings from huddling in a ne.st. Although R. megalotis is active throughout the year, its surface activity is presumably re- duced during the colder months to minimize thermal stress. A male, with a radioactive tag, was monitored for three nights in No- vember (Table 3). The average time spent above ground was 3.3 hours. I accepted this time as representative of both fall and winter activity. This estimate is probably a max- imum because individuals may go several days without any surface activity during in- clement weather. In fact, both Pearson (1960) and Gaertner (1968) allude to hypometabol- ism and possible torpor in Reithrodontomys. In the spring and summer, food availability and a more energetically favorable micro- climate probably extend surface activity. Ac- tivity during these seasons was estimated as 4 hours/night, a value also used by Pearson (1960). Estimated annual energy expenditure for an individual harvest mouse is shown in Table 4. Daily cost for maintenance in males (mean weight 10.76 g) and females (mean weight 9.48 g) is 9.11 and 8.02 kcal, respec- tively. This amounts to 0.85 kcal/g/day, of which thermoregulation accounts for about 74 percent and activity for 26 percent. Maintenance costs were 25 percent lower in Table 2. Mlcroenvironmental temperatures at the Hanford Study Area. Time interval Mean temperature, °C In burrow At soil surface Length Day Night (days) Midpoint (Tb) (Td) (T„) 12 Jun22 24.2 43.9 22.4 36 Jul 16 26.9 40.1 22.5 38 Aug 22 26.0 .37.0 18.5 32 Sep 26 19.2 21.6 8.2 34 Oct 29 12.6 8.6 -1.2 20 Nov 25 lO.©* -1.8b _5jb 27 Dec 19 8.3" -4.5 -7.3 34 Jan 18 5.6* 0.3 -4.5 36 Feb 22 5.6* 5.2 -3.6 30 Mar 27 10.0 18.3 2.4 30 Apr 26 16.5 29.8 11.0 36 May 29 19.7 32.2 15.7 Jun 17-Jun 28 Jun 29-Aug 3 Aug 4-Sep 10 Sep 11-Oct 12 Oct 1.3-Nov 15 Nov 16-Dec 5 Dec 6-Jan 1 Jan 2-Feb 4 Feb 5-Mar 12 Mar 1,3- Apr 11 Apr 12-May 11 May 12-Jun 16 "Temperatures taken at 9 dm (Stone, et al., 1972) "Based on 14 days data 148 Great Basin Naturalist Vol. 39, No. 2 Table 3. Activity patterns and average amount of time spent on the surface by a male Reithrodontomys megalotis. Date Time out time Total time" November 27-28 November 28-29 November 29-30 21:26 1:30 2:43 3:54 4:45 7:08 23:29 1:50 2:54 4:03 5:20 7:33 2h:03m Oh :20m Oh:llm Oh :09m Oh :35m Oh :25m 2h:03m Oh :20m Oh: 11m 0h:09m Oh :35m Oh :18m Total 3h:36m 21:16 22:35 lh:19m lh:01m 23:40 23:56 Oh: 16m Oh: 16m 1:50 2:10 Oh:20m Oh :20m 3:37 3:58 Oh:21m 0h:21m 4:42 4:52 Oh: 10m Oh: 10m 5:47 6:12 Oh :25m 0h:23m Total 2h:31m 20:10 21:07 Oh :57m Oh:57m 22:17 22:42 Oh :25m 0h:25m 0:24 0:50 0h:26m 0h:26m 3:12 4:01 0h:49m Oh :49m 5:32 6:15 Oh :43m 0h:43m 6:47 7:07 0h:20m 0h:20m Total 3h:40m Mean time . . . 3h:16m 'Elapsed time i ; time for periodic retreats to the burrow Table 4. Annual maintenance energy expenditure for Reithrodontomijs megalotis at the Hanford Study Area. Body weight: males = 10.76 g, females 9.48 g. Time interval Hours daily Metabolic costs (kc-ilf Dates Days At rest Active Male Female Er Ea Em Er Ea Em 6/17-6/28 12 20 4 62.0 20.6 82.6 54.6 18.1 72.7 6/29-8/3 36 20 4 159.9 62.5 222.4 140.9 55.0 195.9 8/4-9/10 38 20 4 172.7 73.8 246.5 152.2 65.0 217.2 9/11-10/12 32 20.7 3.3 195.0 66.5 261.5 171.1 58.6 229.7 10/13-11/15 34 20.7 3.3 254.4 84.0 338.4 224.2 74.0 298.2 11/16-12/5 20 20.7 3.3 156,1 52.8 208.9 137.5 46.6 184.1 12/6-1/1 27 20.7 3.3 219.4 74.9 294.3 193.3 66.0 259.3 1/2-2/4 34 20.7 3.3 301.7 89.2 390.9 265.8 78.6 344.4 2/5-3/12 36 20.7 3.3 319.4 92.6 412.0 281.4 81.6 363.0 3/13-4/11 30 20 4 229.3 84.3 313.6 202.0 74.3 276.3 4/12-5/11 30 20 4 195.2 70.6 265.8 172.0 62.2 234.2 5/12-6/16 36 20 4 212.0 75.1 287.1 186.7 66.2 252.9 Annual total (Kcal/yr) without huddling 2477.1 846.9 3324.0 2181.7 746.2 2927.9 (%E„) (74.5) (25.5) (74.5) (25.5) with huddling 2289.1 846.7 3135.8 2016.2 746.2 2762.4 (%E„) (73.0) (27.0) (73.0) (27.0) ^Based on microenvironmental temperatures (Table 2) lune 1979 Sc:hreiber: Rodent Energy Budgets 149 the spring and summer than in the fall and winter. For a 9 g individual of this species in central California, Pearson (1960) estimated daily energy costs averaged 7.6 kcal. His lower value reflects both the smaller average weight and the higher microenvironmental temperatures encountered by that popu- lation. The coefficient of digestibility of this spe- cies, based on the ash tracer method, was 0.856. After adding the energy cost of pre- adult growth (Eg^)^ ingestion rate for a male was calculated as: In, = 3473/0.856 = 4057 kcal/yr (7) Based on a diet of 5.92 kcal/g (mean calor- ic value of stomach material, Schreiber 1979), a male would consume 686 g per year or about 1.9 g per day. Weight gains were determined for each growth stage. At birth harvest mice weigh 1.5 g (Svihla 1931). This was increased 0.4 g to account for embryonic tissues. Weaning weight, prorated from that for deer mice, was calculated as 5.4 g. Mean body weight for adult males and nongravid (NG) females was 10.12 g (N = 34). Caloric density for adult tissue was 1.58 kcal/g (Schreiber and Johnson 1975); other caloric values and growth efficiencies were given in the meth- ods. Mean litter size (n) was 3.6 (N = 8), with females producing 1.11 litters per year (L). The annual ingestion rate for a female was calculated as: If = (2928 + 318)/0.856 = 3791 kcal/yr (8) These mice expend 16.7 percent of this growth energy between conception and birth, 35.7 percent between birth and wean- ing, and 47.6 percent after weaning. Females would annually consume 642 g or 1.8 g per day based on the above diet. Huddling during the colder months would reduce total energy expenditure 5.7 percent. Peromyscus maniculatus: Deer mice were the second most abundant rodent in the study area (Table 1). Because this species is also ac- tive periodically throughout the year, their activity patterns are presumably similar to those determined for harvest mice. For Ep I adjusted the minimum rate measured by McNab and Morrison (1963, Table 1 and Fig. 3) by 24.5 percent, as suggested by Chew and Chew (1970), to obtain the average resting metabolism: E, = 9.3 -0.2 Tb (Tb> 27.1 C) (9) Correcting this for the energy used during activity: The insulating effects of a nest reduce E^ to: E„ = 7.4 -0.13 Tb (Tb<27.1C) (11) Daily, males require about 14.1 kcal and females 12.1 kcal for maintenance (Table 5) or 0.63 kcal/g/day in the spring and summer. Adding the growth increment and using a coefficient of digestibility of 0.879 calculated for animals living in the wild, the ingestion rate for males was: I„, = 5344/0.879 = 6080 kcal/yr (12) In addition to maintenance costs, females in the study areas produced an average of 1.32 litters per year with a mean number of 4.7 young (N = 44). At birth, deer mice weigh an average of 1.8 g (Svihla 1934); embryonic tissues add 1.1 g. Weaning takes place in about 25 days, when animals weigh about 11 g (Svihla 1934, Chew and Chew 1970). Mean adult body weight of males and NG females was 17.5 g (N = 162) and adult tissue of this species has a caloric value of 1.56 kcal/g (Schreiber and Johnson 1975). Previously cited values were used for other tissue energy and growth efficiencies. Ingestion rate for fe- males was: If = (4425 + 753)/0.879 = 5891 kcal/yr (13) Of the 753 kcal used for growth, 16.9 per- cent was used before birth, 56.1 percent from birth to weaning, and 26.9 percent after weaning. Deer mice on the study area con- sumed diets with a mean caloric value of 5.75 kcal/ g (Schreiber 1979). Therefore, to meet the required energy demands, males would consume 1.06 kg/yr and females 1.03 kg/yr, or about 2.9 g/day. This amounts to 14.6 per- cent and 16 percent of the body weight in males and females, respectively. Johnson and Groepper (1970) estimated a 20 g deer mouse in the North Plains consumed 1.9 g of food daily or about 9 percent of its body weight. Deer mice on standard rations at temper- 150 Great Basin Naturalist Vol. 39, No. 2 atures 10-15 C reportedly consumed an aver- age of 3 g/day (Hatfield, 1940) or 2.8 g/day at 28 C (Sealander 1952). Energy requirements effecting reproduc- tive stress may substantially increase (e.g., 200 percent) between birth and the end of weaning (Stebbins 1977), so my estimates for females may be somewhat low. Besides huddling, which effectively reduces metabol- ic costs 5.5 percent (Table 5), deer mice no doubt also resort to additional means of re- ducing energy expenditure during periods of stress. Marten (1973) found this species may sharply reduce its activity through the sum- mer and activity becomes compressed into the early part of the night, when conditions probably permit a more favorable heat ex- change. For the same reason, diurnality may increase in colder months. In winter, pelage (insulation) increases and also contributes to a lower metabolism. Howard (1951) suggested huddling is an important part of energy re- duction in deer mice, with torpor occurring in grouped animals. Torpor in this species has also been observed by others (e.g., Morhardt and Hudson 1966, Morhardt 1970). Addition- ally, Kritzman (1974) and O'Farrell (1975a) suggested possible summer estivation or hy- pothermia for animals at the Hanford Reser- vation. Although the rhythmicity of torpor in natural populations is unknown, it could serve both to conserve water in the summer and reduce energy costs at low temperatures or during periods of food scarcity. Hart (1958) also suggested possible alterations in the ability to metabolize food during such periods of stress. As an additional factor, food caches may offer a significant buffer during periods of severe weather and with sufficient stored food, individuals would spend less time on the surface exposed to unfavorable micro- environmental temperatures. Peromijscus maniculatus is omnivorous, demonstrating marked seasonal variation in its diet (Johnson 1964). On other parts of the reservations this species relied heavily on in- sects for food from spring until fall (Kritzman 1974), probably switching to a more gran- ivorous diet as this food source diminished. It would appear then that competition for available seeds between this species and the more abundant P. parvus would be reduced due to their periods of activity, deer mice be- coming more dependent on seeds in the cold- er months when pocket mice are dormant. Additional research is necessary to properly evaluate these aspects of deer mice energet- ics. Onijchomijs leucogaster: At an average weight of 25 g, the stout-bodied northern grasshopper mouse was the largest rodent species on the study area. It was present in Table 5. Annual maintenance energy expenditure for Peromijscus maniculatus at the Hanford Study Area. Body weight: males = 19.17 g, females 16.83 g. Time interval Hours daily Metabolic costs (kcal)^ Dates Days At rest Active Male Female Er Ea En, Er E, E„, 6/17-6/28 12 20 4 95.0 34.2 129.2 83.4 30.0 113.4 6/29-8/3 36 20 4 258.4 102.0 360.4 226.8 89.6 316.4 8/4-9/10 38 20 4 283.2 118.9 402.1 248.6 104.4 353.0 9/11-10/12 32 20.7 3.3 301.7 102.5 504.2 264.9 90.0 354.9 10/13-11/15 34 20.7 3.3 378.2 128.5 506.7 332.0 112.8 444.8 11/16-12/5 20 20.7 3.3 232.4 80.3 312.7 204.0 70.5 274.5 12/6-1/1 27 20.7 3.3 329.1 111.9 441.0 289.0 98.2 387.2 1/2-2/4 34 20.7 3.3 433.9 135.2 569.1 380.9 118.7 499.6 2/5-3/12 36 20.7 3.3 459.4 141.0 600.4 403.3 123.8 527.1 3/13-4/11 30 20 4 336.8 130.0 466.8 295.7 114.1 409.8 4/12-5/11 30 20 4 292.6 110.4 403.0 256.9 96.9 353.8 5/12-6/16 36 20 4 324.6 120.6 445.2 285.0 105.8 390.8 Annual total (Kcal/yr) without huddling 3825.3 1315.5 5140.8 3270.5 1154.8 4425.3 (%Em) (74.4) (25.6) (73.9) (26.1) with huddling 3547.8 1315.5 4863.3 3026.9 1154.8 4181.7 (%E„,) (73.0) (27.0) (72.4) (27.6) Based on microenvironmental temperatures (Table 2) June 1979 Sc:hreiher: Rodent Enercy Bui)(;ets 151 low numbers throughout the year. Traps holding these mice were often adjacent to those with mutilated pocket mice, indicating O. leiicogastcr's carnivorous tendency and predatory feeding habits. To my knowledge, no metabolic-temperature function equation presently exists for this species, so I used the equation for O. torridus (Chew and Chew 1970), a species of similar size. Average rest- ing metabolism is: Er = 7.24 -0.17 Tb (Tb>27.1C) (14) This species uses a nest (Ruffer 1965) and, assuming Pearson's (1960) correction for its insulating properties, E„ = 5.86 - 2.12 Tb (Tb<27.1 C) (15) above-ground activity increases energy ex- penditure; so, E^ = 10.14- 0.17 Ta (16) presumably, this species has periods of activi- ty similar to harvest mice and deer mice. An- nual maintenance costs were estimated as 4857 and 5215 kcal for males and females, re- spectively (Table 6). Generally, metabolic costs are about 25 percent greater in the colder months than in the warmer months. Nest burrows are shared by male-female pairs (Ruffer 1965); such huddling would produce a 5 percent savings in energy. Males digest an average of 90.3 percent of their caloric diet. E^,^ = 303 kcal for this spe- cies, so ingestion rate was calculated as: I^ = 5160/0.903 = 5714 kcal/yr (without huddling) = 4893/0.903 = 5419 kcal/yr (with huddling) (17) Grasshopper mice weigh 2.2 g at birth (Svihla 1936), excluding embryonic tissues (~0.6 g). Females raised an average of 1.07 litters per year with a mean litter of 3.3 young (N = 14). Young are weaned at about 23 days at an average weight of 13.2 g (Pin- ter 1970) and the mean weight of mature males and NG females was 22.6 g. Using a caloric value of 1.61 kcal/g for adult tissue (Schreiber and Johnson 1975) and other val- ues cited earlier, energy cost for growth from conception to subadult is 660 kcal. The ma- jority of this growth energy is expended after birth. Prenatal growth accounted for 10.6 percent, weaning period for 43.5 percent, and postweaning growth for 45.9 percent. With a digestibility coefficient of 0.892, ingestion rate for females is: If = (5215 + 660)/0.892 = 6587 kcal/yr (without huddling) Table 6. Annual maintenance energy expenditure for Onychomys leucogaster at the Hanford Study Area. Body weight: males = 24.3 g, females 26.2 g. Time interval Hours daily Metabolic costs (Kcal)^ Dates Days At rest Active Male Female Er Ea En, Er E. E„, 6/17-6/28 12 20 4 82.6 35.6 118.2 89.0 38.3 127.3 6/29-8/,3 36 20 4 219.2 105.0 324.2 236.3 113.2 349.5 8/4-9/10 38 20 4 239.3 124.1 363.4 258.0 133.8 371.2 9/11-10/12 32 20.7 3.3 274.3 107.2 381.5 295.7 115.5 411.2 10/13-11/15 34 20.7 3.3 357.1 134.8 491.9 385.0 145.3 5.30.3 11/16-12/5 20 20.7 3.3 222.1 83.9 306.0 239.5 90.5 330.0 12/6-1/1 27 20.7 3.3 316.2 117.4 433.6 340.9 126.6 467.5 1/2-2/4 34 20.7 3.3 426.9 142.6 569.5 460.2 153.8 614.0 2/5-3/12 36 20.7 3.3 452.0 147.6 599.6 487.3 159.1 646.4 3/13-4/11 30 20 4 325.4 135.1 460.5 350.9 145.6 • 495.5 4/12-5/11 30 20 4 272.9 115.5 .388.4 294.3 124.5 418.8 5/12-6/16 36 20 4 293.9 126.0 419.9 316.9 135.8 452.7 Annual total (Kcal/yr) without h uddling .3481.9 1374.8 4856.7 3733.7 1482.0 5215.4 (%E„) (71.7) (28.3) (71.6) (28.4) with h uddling 3215.6 1374.8 4590.4 3446.3 1482.0 4928.3 (%E„,) (70.1) (29.9) (69.9) (30.1) Based on microenvironmental temperatures (Table 2) 152 Great Basin Naturalist Vol. 39, No. 2 (4928 + 660)70.892 = 6265 kcal/yr (with huddling) (18) Males and females which huddle must daily digest 15.6 (0.69) and 17.2 kcal (0.76 kcal/g), respectively. Diets of grasshopper mice include a variety of insects and seeds plus some animal flesh and green vegetation. Stomach material of this species at Hanford had a mean caloric value of 5.22 kcal/g (Schreiber 1979), about 8 percent lower than the value reported for North Plains individ- uals (Johnson and Groepper 1970). Based on this caloric diet each male and female in the population studied would annually consume about 1038 g (2.9) and 1200 g (3.3 g/day) of food, respectively. By comparison, in the lab- oratory on diets of beef liver (72 percent wa- ter), grasshopper mice consumed 4.1 g/day (Whitford and Conley 1971). The incidence of seeds in the diet of this species may significantly increase in the fall and winter (Flake 1971), suggesting it relies on seed caches when insects are less avail- able. Less surface activity would decrease metabolic demands in the colder months. To facilitate this energy savings individuals may restrict more of their winter "hunting" activ- ity to burrow systems. Burrows serve as hi- bernacula for insects, lizards, and torpid mice, all potential food sources for O. leuco- gaster. The generally high fat deposits of this rodent probably conserve body heat when it is forced to be active on the surface during the cold months. Ecological Relationships Individually, P. maniculatiis and O. leuco- gaster process between 2^2 to 3 times as much energy during the year as P. parvus, the most common species (Table 7). The smaller size of the pocket mouse and its peri- odic use of torpor account for this difference. Based on the relative numbers of individuals (Table 1), however, P. parvus dominates as the primary "energy mover" in the small mammal community of this cold desert eco- system. At the population level, the annual contribution of pocket mice to community energy exchange is nearly 4 times that of deer mice, 11 times that of grasshopper mice, and about 17 times that of harvest mice. Even at high population levels, however, the granivorous pocket mouse does not signifi- cantly affect its primary food resource, cheat- grass (Schreiber 1978b). Thus, it is reasonable to assume the euryphagic and less abundant species represented in this paper would also have an insignificant impact on their diverse food resources. These results additionally refine our under- standing of energetics of small mammal com- Table 7. Estimated annual ingestion rates of selected Great Basin rodents. Species Sex (kcal/yr) Coefficient of digesti- bility Mean litter size Number litters per year Energy for' growth (kcal) Ingestion rate (kcal/yr) (kg/yr)b PerognathuS^ parvus M F 2010 1774 0.892 0.911 3.9 1.10 205 360 2483 2342 0.50 0.47 Peromyscus maniculatus M F 5141 4425 0.879 0.879 4.7 1..32 203 753 6080 5891 1.06 1.02 Onychomys leucogaster M F 4857 5215 0.903 0.892 3.3 1.07 303 660 5714 6587 1.09 1.26 Reithrodontomys megalotis M F 3324 2928 0.856 0.856 3.6 1.11 148 318 4057 3791 0.68 0.64 ''Maintenance energy (E^) is calculated without the advantage of savings from hudding. "Based on the mean caloric values of diets of wild mice (Schreiber, 1979) Taken from Schreiber (1978b) June 1979 Schreiber: Rodent Energy Budgets 153 munities. Unfortunately, the question of the ecological significance of these consumers re- mains imanswered and a matter of consid- erable discussion (e.g., Naumov 1975, Chew 1978). As deserts come under increasing pres- sure for development, the understanding of community function and of the relative cost/benefit of perturbing its various com- ponents becomes imperative. Research must now focus on integrating information on bioenergetics with other parameters, such as resource allocation, nutrient cycling and inter- and intraspecific competition, and de- veloping new hypotheses on the role of small mammal consumers. Acknowledgments I am indebted to Dr. Donald R. Johnson, University of Idaho, who initiated this proj- ect and secured its financial support. Addi- tional support and facilities were provided by the Joint Center for Graduate Studies and Battelle Pacific Northwest Laboratories, Richland, Washington (USAEC Contract AT(45-l)-2042). The research represents a portion of my dissertation, submitted in par- tial fulfillment of the requirements for a Doc- tor of Philosophy degree in zoology from the University of Idaho. Final typing was provid- ed by Sally Vreeland. Literature Cited Black, H. L., and N. C. Frischknecht. 1971. Relative abundance of mice on seeded sagebrush-grass range in relation to grazing. U.S.D.A. Forest Serv. Res. Note INT-147. 8 p. Chew, R. M. 1978. The impact of small mammals on ecosystem structure and function, p. 167-180. In: D. P. Snyder (ed.) Populations of small mammals under natural conditions. Spec. Publ. Series, Vol. 5, Pymatuning aboratory of Ecology, Univ. Pitts- burgh, Linesville, Pennsylvania. 2.37 p. Chew, R. M., and A. E. Chew. 1970. Energy relation- ships of the mammals of a desert shrub {Larrea tridentata) community. Ecol. Monogr. 40:1-21. Daubenmire, R. 1970. Steppe vegetation of Washington. Washington State Agri. Exp. Sta. Tech. Bull. 62, Coll. of Agri., Washington State Univ., 131 p. Drozdz, a. a., Gorecki, and K. Sawicka-Kapusta. 1972. Bioenergetics of growth in common voles. Acta Theriol. 17:245-257. Flake, L. 1971. An ecological study of rodents in a short-grass prairie of northeastern Colorado. U.S./IBP, Grasslands Biome Tech. Rep. 100. Col- orado State Univ., Fort Collins. 118 p. French, N. R., B. G. Maza, and A. P. Aschwandern. 1966. Periodicity of desert rodent activity. Sci- ence 154:1194-1195. Gaertner, R. a. 1968. Seasonal variations in the energy budgets of the harvest mouse, Reithrodontomys fiilvescens, and the cotton rat, Sigmodon his- pidus. Unpublished dissertation. Univ. of Ar- kansas. 149 p. Gebczynska, Z., and M. Gebczynskl 1971. Insulating properties of the nest and social temperature regulation in Clethorionomys glareolus (Schreber) Ann Zool. Fennici 8:104-108. Gorecki, A. 1965. Energy values of body in small mam- mals. Acta Theriol. 10:333-352. Grodzinski, W., and a. Goreckl 1967. Daily energy budgets of small rodents, p. 295-314. In: K. Pet- ruscwicz, (ed.), Secondary productivity of terres- trial ecosystems. Panstwowe Wydawnictwo Naukowe, Poland. Hansson, L. 1967. 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Metabolic characteristics of mountain, desert, and coastal populations of Peromijscus. Ecol. 42:723-740. Myrcha, a., and W. Walkowa. 1968. Changes of the caloric value of the body during the postnatal growth development of white mice. Acta Theriol. 13:391-400. Naumov, N. p. 1975. The role of rodents in ecosystems of the northern deserts of Eurasia, p. 299-309. In: F. B. Golley, et al. (eds.). Small mammals: their productivity and population dynamics. Intern. Biol. Prog. 5, Cambridge Univ. Press, Cambridge. 451 p. O'Farrell, T. p. 1975a. Small mammals, their parasites and pathologic lesions on the Arid Lands Ecology Reserve, Benton County, Washington. Amer. Midi. Nat. 93:377-387. 1975b. Seasonal and altitudinal variations in pop- ulations of small mammals on Rattlesnake Moun- tain, Washington. Amer. Midi. Nat. 94:190-204. O'Farrell, T. P., R. J. Olson, R. O. Gilbert, and J. D. Hedlund. 1975. 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Bioenergetics of the Great Basin pocket mouse, Perognathus parvus. Acta Theriol. 23:469-488. 1979. Coefficients of digestibility and caloric diet of rodents in the northern Great Basin desert. J. Mammal. 60: (In press). Schreiber, R. K., and D. R. Johnson. 1975. Seasonal changes in body composition and caloric content of Great Basin rodents. Acta Theriol. 20:343-364. Sealander, J. A., Jr. 1952. Food consumption in Per- omyscus in relation to air temperature and pre- vious thermal experience. J. Mammal. .33:206-218. SoHOLT, L. F. 1973. Consumption of primary productiv- ity by a population of kangaroo rats {Dipodomys merriami) in the Mojave Desert. Ecol. Monogr. 43:357-376. Stebbins, L. L. 1977. Energy requirements during repro- duction of Peromyscus manic iilahis. Can. J. zool. 55:1701-1704. Stone, W. A., D. E. Jenne, and J. M. Thorp. 1972. Cli- matology of the Hanford Area. BNWL-1605, June 1972. Battelle Pacific Northwest Lab., Richland, Washington. SviHLA, A. 19.34. Development and growth of deer mice (Peromyscus manlculatus artemisiae). J. Mammal. 15:99-104. SviHLA, R. D. 1931. Notes on desert and dusky harvest mice (Reithrodontomys megalotis megalotis and Rcithrodontomys megalotis nigrescens. J. Mam- mal. 12:363-365. 1936. Breeding and young of the grasshopper mouse (Onychomys leticogaster fuscogriseus). J. Mammal. 17:172-173. Trojan, P., and B. Wojciechowska. 1968. The effects of huddling on resting metabolism rate in Eu- ropean common vole Microttis arvalis. Bull. Acad. Pol. Sci. CI. II. 16:107-109. Whitford, W. G., and M. I. Conley. 1971. Oxygen consumption and water metabolism in a carni- vorous mouse. Comp. Biochem. Physiol. 40:797-803. Wiener, J. G., and M. H. Smith. 1972. Relative effi- ciencies of four small mammal traps. J. Mammal. 53:868-873. i PRELIMINARY SURVEY OF RAPTOR SPECIES IN THE MANTI DIVISION, MANTI-LA SAL NATIONAL FOREST Stephen C. Jones' Abstract.— A preliminary survey of raptor species and habitat in the Manti Division, Manti-LaSal National For- est, was condvicted during May 1977. Eight species of raptors were recorded, the most common being the Golden Eagle {Aquila chnjsaetos) and American Kestrel {Falco sparveritts). Nine additional nesting and migratory species were reported by cooperating observers and based on habitat analysis another two species are expected. Eleven Golden Eagles, representing at least eight pairs, were observed and it is estimated that from 15 to 20 pairs of eagles inhabit the division. The Manti Division provides habitat for two endangered species, the Peregrine Falcon (Falco peregrinus anatttm) and the Northern Bald Eagle [Haliaeetus leucocephahis alascanus). Further research is needed to ascertain the impact that current development activities in the division will have on the raptor community. With the increased awareness of the criti- cal role that raptors play in balanced ecosys- tems and the declining nature of many spe- cies, land management for the conservation of birds of prey has become an important consideration of governmental agencies ad- ministering public lands (White 1974, Olendorff and Kochert 1975). Although U.S. Forest Service lands are administered under the multiple land use philosophy (Griswold, 1978) the conservation of habitat and the protection of areas critical to threatened or endangered species are of prime importance. Paramount to this goal is the identification of species inhabiting an area and assessment of available habitat for species suitability. The present study was conducted to identify the raptor species inhabiting the Manti Division, Manti-LaSal National Forest, and to assess the available habitat for suitability to raptors. This survey takes on added significance be- cause this area is being impacted by the con- struction and operation of energy-operating facilities. Judgments on critical habitat and the impact of human activity in the division on the raptor community were made but are presented elsewhere (Jones 1977). . Study Area and Methods The Manti Division is located in central Utah about 32 km west of Price and extends southward for approximately 120 km (Fig. 1). Elevation ranges from 1,400 to 3,500 m above sea level. The east rim of the Wasatch Plateau forms the eastern boundary of the di- vision. The predominant vegetation types are pinyon-juniper and sagebrush and, at higher elevations, aspen and spruce-fir. Approx- imately 8 percent of the division is under pri- vate ownership. Seven coal mines and four sawmills are operating in or near the national forest. This study was conducted in the south- eastern portion of the Manti Division, en- compassing an area of about 960 sq. km. The major canyon systems, the intermittant and permanent streams, and the associated ripa- rian areas provide the habitat used by raptors in the division and were the focal points of this study. Field work was conducted from 1 May to 15 May 1977. Each of the canyon systems and areas shown in Figure 1 was examined at least once, and in most cases twice. Vehicle travel was possible in all areas except Upper Rocky and Muddy Creek canyons. Raptors were observed and cliff areas examined with 7 X 35 binoculars and a 20X spotting tele- scope. Data were collected on species seen, nests located, and the suitability of the avail- able cliff, riparian, prey, and roosting habi- tats present. All nests were delineated on uses IVi minute topographic maps for use by Forest Service personnel. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 155 156 Great Basin Naturalist Vol. 39, No. 2 Fig. 1. Mant. Div.ion, Manti-LaSa. National Forest. N.u.bered areas f ^ ^ -J°-y;rr"T^eXrS^^^^^^^ for rfptor species and habitat: ^ Huntington ^^^^^- ^:^::, ^.w n 8. Stt^tlnyon, 9. Lower Joes 5. Crandall Canyon, 6. Grimes Wash and Danish Bam 7. Cotton^^^^^^^^ ^ Upper Rock Canvon, 1.3. Lower ^C:^Z&l^^::;'fr^:^ - -on Snyon overlook and Flagsta. Peak, 16. Muddy Creek Canyon. / June 1979 Jones: Raptoh Survey 157 The analysis of habitat suitability was based on actual sightings or the probability of use by raptors. Judgments on the probable use of an area were made according to the habitat preferences of each raptor species as described in McGahan (1968), Camenzind (1969), and Beecham and Kochert (1975) for the Golden Eagle [AquUa chrysaetos); Porter and White (1974) for the Prairie Falcon (Fal- co mexicanus) and Peregrine Falcon (F. per- egrinus anatum); Smith and Murphy (1973) and Hay ward et al. (1976) for hawks and owls in Utah; Jones (1978) for Accipiter hawks and Joseph (1977) for the Northern Bald Eagle {Haliaeetus leucocephalus alas- canus) wintering in Utah. Judgments about prey habitat were made based on numbers and diversity of potential prey species seen. Special efforts were made to locate eagle nests, as the Golden Eagle is the most sensi- tive raptor to habitat destruction and disturb- ance nesting in the division. Results Table 1 presents a list of the eight species of raptor seen and an analysis of the habitat suitability for each of the areas visited. Most commonly seen were Golden Eagles and the American Kestrel (F. sparverius), with the Red-tailed Hawk {Buteo jamaicensis) and Cooper's Hawk {Accipiter cooperii) being seen on several occasions and a Goshawk (A. gentilis). Sharp-shinned Hawk (A. striatus), and Prairie Falcon each observed once. Great-horned Owl {Bubo virginianns) cavities were seen in five areas. In addition to my sightings, cooperating observers recorded other species of raptors for the division. C. Jemmett, U.S. Forest Ser- vice, reports (pers. comm.) occasional Osprey {Pandion haliaetus), wintering Bald Eagles, and a Peregrine Falcon in the division. The Ferruginous Hawk {B. regalis), Swainson's Hawk {B. swainsoni), Turkey Vulture {Cath- Table 1. Raptor species seen and habitat suitability in Manti Division, Manti-LaSal National Forest. Areas surveyed Species observed and expected' Habitat' s c3 8 -g 2 o -5b 2 1 1 * s. o O ^ u (C O w Oh 1^ o u B cu tf 1. Huntington C. + - p p - - - + - L,I I - 2. RildaC. P P - - - - - - - I I - 3. Mill Fork C. P P _ p + p - - - I,H I - 4. Tie Fork C. P P _ p - - _ - _ 1 I - 5. Crandall C. -1- + + - - - - - - L H - 6. Grimes Wash, Danish Basin - - - - + p - P - L,l - H - 7. Cottonwood C. P P - p + + - + + I,H 1,H I,H - 8. Straight C. + - + - + + - + + I,H I,H I - 9. Lower Joes Valley, Lowry C. - + - + -1- + + + + LH I,H H H 10. Upper Joes Valley P - - p + p - P - - I H - 11. North Horn Mtn. - - - - + - - + - - - H - 12. Upper Rock C. - - - - + + - P - I,H - L,I ■ - 1.3. Lower Rock C, East Rim - - - - + - - - + L,I - L,I - 14. Ferron C, Bull Hollow - - P _ + p - + - L,I I,H LH H 15. Ferron C, Flagstaff Peak P _ _ + - _ - P + L,l - I - 16. Muddy Creek C. - - + - - - - P - L,I I I - 1. Species and structures: + = Observed, - = Not observed, P = Probable occurrence. 2. Cavities assumed used by Great-homed Owls. 3. Habitat suitability: L = Low, I = Intermediate, H = High 4. Roosting habitat suitability judged with respect to wintering Bald Eagles. 158 Great Basin Naturalist Vol. 39, No. 2 artes aura). Marsh Hawk {Circus cyaneus). Saw-whet Owl (Aegolius acadicus), and Flammulated Owl {Otm flammeohis) have been recorded by Phil Wagner, Division of Wildlife Resources (pers. comm.). However, several of these species inhabit the division only during migration. Based on habitat anal- ysis and distribution accounts in Behle and Perry (1975) and Hay ward et al. (1976), an- other two species, the Screech Owl (O. asio) and Pygmy Owl {Glaucidium gnoina), should also occur in the Manti Division. Eleven Golden Eagles, representing at least eight pairs, were seen within nine areas. The eagle sighted on North Horn Mountain is believed to nest in Upper Rock Canyon. Sev- en eagle nests were found within four areas; centers of these nesting areas were from 6.4 to 12.8 km apart. Approximate home range sizes for the eagles in the division can thus be calculated as 44 to 132 sq. km, assuming a roughly circular territory (see Smith and Murphy 1973). Goshawks, Sharp-shinned, and Cooper's hawks appear to be rather common residents in the Manti Division. Suitable riparian habi- tat in most areas was occupied or suspected to be occupied by at least one of the three species. Typically, the Cooper's Hawk will select the lower elevations with more open habitat, with the Goshawk and Sharp-shinned Hawk nesting higher up and accepting steep canyons. Crandall Canyon supported all three species. Due to the secretive nature of these hawks, observations are uncommon even in areas where they are nesting. Red-tailed Hawks should occur more com- monly in the division than the few actual sightings would indicate. They are suspected of nesting on many of the same cliffs that the eagles were using. During winter the Red- tailed Hawk is the most commonly seen hawk in the division (P. Wagner, pers. comm.). The sighting of the Prairie Falcon in Lower Joes Valley is the first record of this species for the division (C. Jemmett, pers. comm.). Suitable nesting areas for Prairie Falcons were found in Lower Joes Valley and Cottonwood Canyon. The individual sighted is believed to nest in the canyons west of Lower Joes Valley. Where seen, Kestrels were numerous. In Lower Joes Valley at least six pairs showed nesting activity during this study. The only habitats not supporting Kestrels were the dry sagebrush areas and the steep mountain can- yons. Cavities suspected of being used by Great- horned Owls were seen in five different areas. Great-horned Owls should be widely distributed in the Manti Division. Nesting cavities do not always show signs of usage, and crow or hawk nests are often used by owls. The other species of owls recorded or suspected are also thought to be widely dis- tributed in the division. A correlation from Table 1 of species pres- ence and habitat availability is indicative of the rather restricted habitat preference of Accipiter hawks and the general preference of eagles, Red-tailed Hawks, and Kestrels. The dry mountainous habitat of the Manti Division will support a certain assemblage of breeding raptors, but other species will be re- stricted. Ultimately, habitat availability and climatic regimes are responsible for the avian species living in an area. In the Manti Divi- sion steep-sided canyons occupied by Accipi- ter hawks open into sloping sagebrush valleys or high aspen meadows that are utilized by eagles, Red-tailed Hawks, and Kestrels. Little falcon habitat is available, except in Lower Joes Valley. The higher elevations restrict such lower desert species as the Ferruginous Hawk. Discussion The Manti Division supports a high diver- sity of raptors, with as many as 17 species utilizing the division during the breeding sea- son and on migration. This study provides an estimation of the breeding raptors within the area surveyed. Wagner (unpubl. ms.) pro- vides data on the raptors utilizing the divi- sion during migration. On 1-2 September 1977 a count of migrating raptors along the Manti Mountain-Skyline Drive yielded the following numbers for each species: Turkey Vulture— 5, Goshawk— 1, Cooper's Hawk— 3, Sharp-shinned Hawk— 1, Red- tailed Hawk— 66, Swainson's Hawk— 1, Ferruginous Hawk— 1, Golden Eagle-4, Kestrel-43, Prairie Fal- con-2 and Marsh Hawk-3 (total = 130). This list is similar to the breeding list with 1979 Jones: Raptor Survey 159 the exception of the relative numbers of each species. Wintering Bald Eagles will utilize Lower Joes Valley reservoir and should also be pres- ent in Perron Canyon. C. Jemmett reports (pers. conim.) that Bald Eagles stay at Lower Joes Valley reservoir as long as waterfowl are present on the reservoir. In severe winters the reservoir and stream freeze over by Janu- ary and are not used by wintering eagles. Im- portant characteristics of areas used by win- tering Bald Eagles are permanent streams which do not freeze over, riparian type vege- tation with a high degree of exposure, and the presence of migrating waterfowl, rabbits, or an abundance of local fishes as prey spe- cies (Joseph 1977). The density of nesting Golden Eagles found on the Manti Division is similar to that found in other western states. McGahan (1968) found the maximum area used by eagles in Montana was from 180 to 205 sq. km. Dixon (1937) reports the area used by 27 nesting pairs in the California foothills ranged from 52 to 160 sq. km. Assuming that eagles are distributed evenly over the entire Manti Division and using the value of 80 sq. km to represent their approximate habitat size requirements, the division should support from 15 to 20 pairs. Considering the number of eagles observed during this rather limited study, the above estimate seems conservative. Nesting densities of eagles in central Utah can, however, vary greatly between years, with cyclic fluctuations in prey densities (Camenzind 1969, Murphy 1975). Analysis of habitat suitability for raptors shows that relatively few areas in the division provide habitat for all species, or complete habitat for any one species (Table 1). The no- table exceptions to this are Crandall Canyon, Cottonwood Canyon, Lower Joes Valley, and Ferron Canyon; these areas provide both nesting and foraging habitat for three to six species. More typically, however, suitable nesting habitat is found in one area and fo- raging habitat in another area. For example. North Horn Mountain may be used for forag- ing by as many as three pairs of Golden Eagles, but on the mountain proper there is no eagle nesting habitat. In addition to the utilization of the habitat within the division by an endangered species. the Bald Eagle, and a sighting of a second en- dangered species, the Peregrine Falcon, sev- eral of the other resident raptors, are expe- riencing population declines in other areas of the United States. The need for protection of populations of these species is, therefore, crit- ical. Important in this is an understanding of the ecology of each species of raptor in- volved, including its breeding requirements, nesting habits, and foraging and food habits. Further research on the raptors in the Manti Division is recommended so that current de- velopment activities do not threaten the spe- cies now utilizing the area. Murphy (1978) outlines management techniques for the western raptors which would be applicable for the Manti Division. Schamberger and Earner (1978) discuss the process of using habitat evaluations in project planning, which is also applicable. The raptors in the Manti Division represent a valuable natural resource, as important for ecological, scien- tific, and aesthetic reasons as energy re- sources and recreational potential are for economic reasons. Acknowledgments Support for this study was provided by the Manti-LaSal National Forest, Contract Q-77- 4. Mr. Coy Jemmett, wildlife biologist, Man- ti-LaSal National Forest, Price, Utah, organ- ized the study and provided administrative assistance. I wish to acknowledge Anita Jor- gensen for her assistance during field work. Literature Cited Beecham, J. J., AND M. N. KocHERT. 1975. Breeding bi- ology of the golden eagle in southwestern Idaho. Wilson Bull. 84:506-513. Behle, W. H., and M. L. Perry. 1975. Utah birds: sea- sonal and ecological occurrence chart and guide to bird finding. Utah Mus. Nat. Hist., Univ. of Utah. 143 pp. Camenzind, F. J. 1969. Nesting ecology and behavior of the golden eagle. In J. R. Murphy et al.. Nesting ecology of raptorial birds in central Utah. Brig- ham Young Univ. Sci. Bull., Biol. Ser. 10(4):4-15. Griswold, R. K. 1978. Forest Service views. Transac- tions of the Forty-third North American Wildlife and Natural Resources Conference. Wildlife Manage. Inst., Wash., pp. 368-371. Hayward, C. L., C. Cottan, A. M. Woodbury, and H. H. Frost. 1976. Birds of Utah. Great Basin Nat. Memoirs 1:1-229. 160 Great Basin Naturalist Vol. 39, No. 2 Jones, S. G. 1977. Raptor species and habitat in the Manti Division, Manti-LaSal National Forest. Unpubl. Final Report. U.S. Forest Service, Man- ti-LaSal National Forest, Price, Utah. 1978. The North American accipiters— goshawk, cooper's hawk, and sharp-shinned hawk. Techni- cal Note No. 17, Bureau of Land Management, In press. Joseph, R. A. 1977. Behavior and age class structure of wintering northern bald eagles {Haliaeetus leii- cocephalus alascanus) in western Utah. Unpubl. M.S. thesis. Brigham Young University. McGahan, J. 1968. Ecology of the golden eagle. Auk 85:1-12. Murphy, J. R. 1975. Status of a golden eagle population in central Utah, 1967-1973. In: Population Status of Raptors, Raptor Research Report No. 3, pp. 91-96, Raptor Research Foundation, South Da- kota. 1978. Management considerations for some west- ern hawks. Transactions of the Forty-Third North American Wildlife and Natural Resources Con- ferences. Wildlife Manage. Inst., Wash., pp. 241-251. Olendorff, R. R., and M. N. Kochert. 1975. Land Management for the conservation of birds of prey. Proceedings, World Conference on Birds of Prey, Vienna, pp. 294-306. Porter, R. D., and C. M. White, in collaboration WITH R. J. Erwin. 1973. The peregrine falcon in Utah, emphasizing ecology and competition with the prairie falcon. Brigham Young Univ. Sci. Bull., Biol. Ser. 18(1): 1-74. Schamberger, M., and a. Farner. 1978. The habitat evaluation procedures: their application in proj- ect planning. Transactions of the Forty-Third North American Wildlife and Natural Resources Conference. Wildlife Manage. Inst., Wash., pp. 274-283. Smith, D. G., and J. R. Murphy. 1973. Breeding ecology of raptors in the eastern Great Basin of Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 18(.3):l-76. White, C. M. 1974. Current problems and techniques in raptor management and conservation. Transac- tions of the Thirty-Ninth North American Wild- life and Natural Resources Conference. Wildlife Manage. Inst., Wash., pp. 301-312. VEGETATION RESPONSE TO A MOISTURE GRADIENT ON AN EPHEMERAL STREAM IN CENTRAL ARIZONA Deborah Ann Bloss' and Jack D. Brotherson- Abstract.— Ecological aspects of desert vegetation in relation to a moisture gradient along an ephemeral stream in central Arizona were investigated. The stream channel, flood plain, and north-, west-, south-, east-facing slopes represent a moisture gradient going from mesic to xeric conditions. Vegetation in some areas of the stream channel intergraded into flood plain vegetation, which in turn intergraded into slope vegetation types. In other areas there were sharp delineations between stream channel and flood plain and between flood plain and slope. Trees and le- giuiie species preferred midmoisture habitats, but forbs, shnibs, and succulents preferred dryer areas. Family groups like the Asteraceae and the Poaceae were found to be distributed ubiquitously. Niche widths were broadest for flood plain species. Diversity was highest on the slopes. Negative correlations existed between stand diversity and the Synthetic Stand Moisture Index (i.e., as moisture increased diversity increased). It is believed that disturbance as well as moisture influenced diversity- Central and southern Arizona are consid- ered to be part of the Sonoran Desert, which occupies much of the southwestern United States and northern Mexico. Phys- iographically, the area is characterized by small mountain ranges rising above level ba- sins and by a vegetation comprised of shrubs, small trees, cacti, and associated ephemerals. The level basins are now liberally marked by arroyos or wadis (i.e., dry washes cut by the action of ephemeral streams). Bryan (1928) suggests that the arroyo cutting is of recent origin. The rainfall pattern of the Sonoran Desert exhibits two distinct peaks: (1) a summer rainy period from July to September charac- terized by thunder storms of short duration and heavy rainfall intensity, and (2) the gentle rains of December through March caused by the southward movement of sub- tropical high pressure systems. Summer rains which are of greater intensity vary less from year to year than do the winter rains (Mal- lery 1936a). Localized storms in the region are very pronounced, often being limited to a few square miles. Temperatures in the Sonoran Desert are characterized by great diurnal fluctuations, hot summers and warm winters. Freezing temperatures may occur, but without regu- larity and they do not last for extended peri- ods of time. Vegetational descriptions of the Sonoran Desert in part at least include Nichol (1952), Whittaker and Niering (1964, 1965), Marks (1950), Glendening and Paulsen (1955), Little (1950), Kramer (1962), Richards (1925), and Shreve (1922, 1924). Spalding (1910), Blumer (1909, 1910) and Walmo (1955) briefly men- tion desert vegetation in foothill sites they studied in the southern Arizona mountain ranges. Keil (1970) and Letho (1970) recently completed floristic studies in two areas of the Sonoran Desert located in Maricopa County. Ecological studies dealing with isolated fragments of Sonoran Desert ecosystems are more numerous than are those dealing with community dynamics or entire community structure. For example, germination studies of annuals (Klikoff 1966, Capon and Asdell 1967, Beatley 1967, Went 1942, 1948, and 1949, Went and Westgaard 1949, Juhren et al. 1956, and Trevis 1958a, 1958b) and of perennials (Turner 1963, Shreve 1931, Turn- er et al. 1966, and Barbour 1968) have been done. Productivity studies of perennials (Chew and Chew 1965) and their ecology in relation to soil types, mineral nutrients, water 'Biology Department, Gila Bend High School, Gila Bend Arizona 85337. 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 161 162 Great Basin Naturalist Vol. 39, No. 2 relations, and plant associations (Yang 1950, Livington 1910, Yang 1957, Gardener 1951, 1959, Spalding 1909, Yang and Lowe 1956, Rich 1961, Reynolds 1962, Muller and Muller 1956, Dalton 1962, Wilm 1956, Parker and Martin 1952, Reynolds and Tshirley 1957, and Halvorson and Patten 1974) have also at- tracted interest. Work with climatic factors in desert areas has been completed by Smith (1956), Sellers (1960), Humphrey (1933), and Mallery (1936a, 1936b). Niering et al. (1963), Turner (1963), and Halvorson (1970) have briefly discussed the pattern of vegetational change from mountain slopes down onto the basin floors. However, their discussions are limited to population dynamics and vegeta- tion studies of foothills and mountain slopes. Halvorson (1970) indicates that the valleys and mountains of central Arizona have not been studied in detail as is the case with the mountain and foothill areas in the more southern portions of the state (see Whittaker and Niering 1964, 1965, Niering et al. 1963, Spalding 1909, 1910). The purpose of this study was to describe the vegetational zona- tion patterns existent along an ephemeral stream in central Arizona and to determine whether the observable vegatational patterns represented a continuum adapted to a mois- ture gradient from slope to stream channel or if they represented distinct, discrete commu- nities. Study Area The study area lies in a foothill region of the Sonoran Desert and is part of the Mexi- can Highlands, a subdivision of the Basin and Range Province, which is geographically lo- cated to the west and south of the Colorado Plateau (Halvorson 1970). The area of study is a five-mile section of New River, an ephemeral stream located 35 miles north of Phoenix, Arizona (Fig. 1). This section of the river parallels Interstate 17 for 2.5 miles and then bends away following Table Mountain and paralleling Table Mesa Road (Fig. 1). New River is a tributary of the Fig. 1. Map of the study area showing location in Arizona. June 1979 Bloss, Bhothkhson: Moisti'HE C^radient 163 Agua Fria River, which feeds into the Gila river and drains most of Central Arizona. Metamorphic, igneous, and sedimentary rock comprise the parent material through which the Gila River and its tributaries flow. Davis (1925) proposed the origin of the exist- ing mountain ranges to be tilted and moder- ately dissected fault blocks. From such ranges the broad desert basins have derived their soil, collected from alluvial material eroded from adjacent mountain slopes. The soils in the study area are generally shallow and poorly developed. Desert pavement is a prominent feature in the area and, as is the case in many desert areas, a narrow surface horizon exists in association with a caliche near the soil surface. The area has been sub- jected to grazing by domestic livestock since the 1800s. Location 2 of the study area is in close proximity to the site of an old stage sta- tion. Much of the area is now controlled by the Tee Cattle Company. Methods Ten study sites were located at one-half- mile intervals along a five-mile section of New River (Fig. 1). Five stands were selected at each site, one stand each at the following locations: both slopes adjacent to the stream channel, the flood plain on both sides of the stream channel, and the stream channel itself. In each stand, three 5 X 10 m plots were set up approximately 3 m apart. Four quadrats, each 1 m square, were placed inside each large plot. Trees and treelike vegetation were sampled within the large plots, and small shrubs, forbs, and grasses were sampled using the meter square quadrats. Frequency data were taken and used in the analysis. Vegetation patterns were analyzed through the use of a synthetic moisture gradient. The moisture gradient was established through knowledge gained from the literature (Nier- ing et al. 1963) and from work done in the field. The stream channel was selected as being the most moist. The two flood plain imits were lumped together and considered the next most moist. In most north temperate floristic systems, the north-facing slopes are considered to be more moist than the oppo- site south-facing slopes (Daubinmire 1974). Such considerations are generally attributed to a shadowing effect of the hills and moun- tains that prevent direct sunlight from ever reaching the area. A similar relationship has been noted with east- and west-facing slopes. However, because of the peculiar shadowing effect of the adjacent hills noted in the study area, it was felt that the east-facing slope would exhibit the dryest conditions, with the west-facing slope fitting in between the north- and south-facing slopes on the mois- ture gradient. Those stands common to each area (i.e., stream channel, floodplain, etc.) were lumped together and average frequency values for each species in the area were com- puted. The species were then arranged in or- der of their occurrence in the six areas (Table 1). Those species most frequent in the stream channel were segregated to the top of the table and those most frequent on the east-fac- ing slope were placed near the bottom, with all others falling somewhere in between. From this table indicator species were se- lected using as a basis frequency values ap- proximately twice that in one group as in any other. The indicator species were then weighted with an index number. The index number was determined as follows: the stream channel species, selected as being rep- resentative of the most moist areas, were as- signed the index number 5; the flood plain species in the next most moist area were as- signed the number 4; and so on with the spe- cies of the east-facing slope receiving the number 0. When the frequency value of a species was not twice that of the next highest group, yet was greater and formed the apex of an increased upward trend, the species was assigned an index number midway between those for the groups in which the two largest frequencies were found and included as an indicator. Synthetic Stand Moisture Index values (SSMI), similar to Plot Index Values (PIV) de- scribed by Dix and Butler (1960), were con- structed by dividing the sum of the compos- ite indices for all indicator species found in a stand by the sum of the stand frequencies of those indicator species found in that stand. The composite index values were determined by multiplying the stand frequency of an in- dicator species by its moisture index number. Therefore, 164 Great Basin Naturalist Vol. 39, No. 2 SSMI 2 composite index values for indicator species 2 frequency values for indicator species SSMI values were computed for all stands. The stands were then placed along a one-di- mensional ordination in an attempt to relate the stands spatially. Indicator species and other vegetational parameters were then graphed against the ordination in an attempt to elucidate trends. Average frequency values were computed for all stands falling between the SSMI values of 0 to 1, 1 to 2, 2 to 3, 3 to 4, 4 to 5, on the moisture gradient. This method was employed because it showed trends much more clearly than the entering of all points. Cluster analysis techniques (Sneath and Sokal 1963) were applied to similarity index figures obtained by utilizing Sorenson's index of similarity (Sorensen 1948) and the follow- ing equation from Dix and Butler (1960): 2W K = a -t- b X 100 Table 1. Average percent frequency values for species distributed in the six groups used to establish the synthetic stand moisture gradient. Numbers to the left of the species names represent index numbers assigned to those species selected as indicator species of the different groups. Species Average percent frequency River Flood- plain Slop e bottom North West South East 83 33 0 0 0 0 55 33 0 0 0 0 45.5 .35.4 0 0 0 0 33 2 0 0 0 0 0 25 14.6 0 0 0 0 19.5 8.3 0 0 0 0 16.7 1.5.9 12.5 0 0 8.3 10.4 8.3 0 0 0 0 8.3 8.3 0 0 0 0 0 8.3 0 0 0 8.3 0 8.3 0 8.3 0 8.3 0 8.3 0 0 0 0 0 8.3 8.3 8.3 0 0 8.3 13.9 8.3 0 0 0 0 19.8 0 0 0 0 16.5 25.2 0 0 0 0 0 33 0 0 0 0 0 53.7 33 44.3 33 33 0 62.2 0 66.7 0 0 33 76.1 49.5 66.7 58 61 0 55 69.8 58.5 .33.2 66.2 0 49.6 66.5 46.5 58 58 13.8 37 56.2 19.4 30.2 41.5 0 8.3 33 37.1 8.3 8.3 0 0 38.8 33.3 8.3 0 0 43.3 49.5 49.5 0 51.4 0 0 0 0 25 8.3 0 26.1 16.7 32 38.9 33.2 0 0 0 8.3 58.3 16.7 0 16.7 0 26.8 66.7 20.6 0 33 55.5 66 82.5 52.1 0 0 55.6 66 1(K) 0 0 0 0 0 0 16.6 0 8.3 0 8.3 16.7 24.9 0 40.3 0 33 .33 49.5 0 33 0 33 33 66.5 0 30.1 8.3 59.7 56.5 80.0 5 Baccharis glutinosa 4.5 Baccharis sarothroides 4.5 Hymenoclea monogtjra 5 Salix sp. 4.5 Pohjpogon monspeliensis 5 Eiiphobia sp. Bromus rubens 4.5 Minulus glabratus 4.5 Datura meteloides Marrubium vulgare Mam»iilhiria microcarpa 4.5 Avciui fatua 3 Ephedra viridis Lepidium sp. 4 Franseria ambrosioides 4.5 Cijnodon dactijlon 4 Phitaiui.s urightii Lyciuin fremontii Acacia greggii Prosopis juli flora Simmondsia chinensis Cerciditim tnicrophyUum Schismus barbatus 2.5 Ferocactus wislizeni 2.5 Eriogonum fasciculatum Larrea divaricate 1 Eriogonum sp. Plantago purshii 1 Opuntia leptocaulis Opuntia engelmannii Opuntia bigclovii 1.5 FoiKjiiicria splendens ^uhohi kali 0.5 Enchinocereus engelmanii Carnegiea gigantea 0 Opuntia fidgida 1 Franseria deltoidea liiiie 1979 Bloss, Brotmerson: Moisture Gradient 165 where "K" ecjuals the index of similarity, "a" is the frequency of all species in one stand, "b" is a similar figure for a second stand, and "W" is that part held in common by both stands. Analyses of the similarity index values and the cluster techniques were carried out on an IBM 7030 computer. Niche width and stand diversity values were computed using the equation B = l/!Spi2 where "B" is equal to either the niche width and/or diversity and "pi" is a measure of the relative abundance of a species in a given habitat (Levins 1966, MacArthur 1972). Niche width values for a species were ob- tained by siunming its pi values across all stands. Diversity values, on the other hand, were computed by summing the pi values for all species found, in a single stand. The P X F (Presence X Frequency) index (Anderson 1964, Curtis 1959) was computed for all species found in the study area to give an indication as to species importance in the system (Table 4). Diversity Indices (DI) were compared to the SSMI by regression analysis (Hall 1971, Dick 1971) to determine if any relationship existed between the two measures. Species determinations follow Kearney and Peebles (1951). Results and Discussion The five-mile section of New River pre- sents a study in contrasts. Characteristically the flood plain tends to be wide in some areas and narrow in others. The vegetation along the wide areas intergrades readily with that of adjacent slopes, but sharp lines of de- marcation appear between flood plain and slope vegetation where the flood plain nar- rows. The flood plain exhibits a wide variety of vegetational pattern. The broader areas display vegetation types closely allied to those of adjacent slopes, with the narrow areas exhibiting greater densities of trees, shrubs, and thicket-forming plants. The river channel itself is wide, sparsely vegetated, and disturbed periodically by high water flow re- sulting from heavy rains in the mountains, evidenced by the highly sorted sand, gravel, and rocky nature of the channel substrate. Seep areas in the channel are characterized by the presence of water nearly year-round. In these areas larger trees such as Salix spp. and Platanus wrightii are foimd along with Mimuhis glahratus and Baccharis guhitinosa. The north-, west-, south-, and east-facing slopes in that sequence appear to be in order of decreasing moisture. Differences in ground cover, the number, and types of individuals on the slopes is apparent. The north-facing slopes generally exhibit a larger more meso- phytic flora. Vegetation analysis involved establishing a synthetic moisture gradient. This was accom- plished by arranging species in order of mois- ture delineations and assigning index numbers (Table 1). Synthetic Stand Moisture Index values (SSMI) were then computed for all stands (Table 2). Using these values the stands were placed on a one-dimensional ordination along the proposed moisture gradient, which oriented them spatially to each other (Fig. 2). Average frequency for indicator species and life form groups in all stands were then graphed against the one-dimensional ordina- tion. Species and life form groups were found to peak in importance along sections of the gradient (Figs. 3-4). Franseria deltoidea, for example, is well adapted to the dryest mois- ture regime on the gradient, but Baccharis sarothroides occurs in more moist sites. Fig- ure 3 also depicts the relationships of several of the indicator species (i.e. Fouqiiieria splen- dens, Ferocactus ivislizeni, and Cercidium mi- crophylhim), for those stands of the study area occurring between 1 and 2 on the mois- ture gradient. Other species exhibiting sim- ilar patterns to those in Figure 3 were Optintia leptocaulis, O. bigelovii, O. engel- mannii, and Plantago purshii (Table 1). Also shown are the patterns exhibited by index plants (i.e., Franseria deltoidea, Opuntia ful- gida, etc.) of the gradient class, 0 to 1. Spe- cies showing similar patterns were Salsola kali and Carnegiea gigantea. The species Hijmenoclea monogijra and Cijnodon dacty- lon showed peak prefence in the most' moist areas (Fig. 3). Baccharis glutinosa, Salix spp.. Euphorbia spp., Mimulus glahratus, and Da- tiire meteloides are other species found to ex- hibit similar patterns. Franseria ambrosioides (Fig. 3) peaked between the SSMI values of 3 and 4, which represents the flood plain area 166 Great Basin Naturalist Vol. 39, No. 2 of the study. Other species prefering this same habitat were Acacia greggii and Sim- mondsia chinensis. It should be noted, how- ever, that these two species are rather ubiqui- tous and may be prominent in other areas as well. Schismtis barbatiis and Eriogonum fas- ciciilatum, though not clear-cut index plants of the west and south slopes (there were no clear-cut indicators for this section of the gradient), show optimum frequency peaks in this area. Because the relationship between various Table 2. Stands listed along with their Synthetic Stand Moisture Index values (SSMI), diversity indicies, aspect and/or topographic position on the landscape. EF = east facing slope, SF= south facing slope, WF = west facing slope, NF = north facing slope, FP = floop plain, and SC = stream channel. Diversity Diversity Index Stand No. SSMI Location" Index Stand No. SSMi Location 1 1.0 EF 5.886 25 4.5 RB .3.019 2 1.0 EF 4.970 26 4.5 RB 2.062 3 .91 Ef 5.394 27 0 RB 0 4 0 EF 4.003 28 4.6 RB 2.538 5 .50 EF 6.270 29 4.7 RB .3.019 6 .83 EF 9.452 30 4.6 RB 3..331 7 1.0 SF 5.388 31 4.5 FP 5.160 8 1.3 SF 5.504 32 4.2 FP 1.790 9 1.0 SF 6.515 33 .90 FP 7.457 10 .67 SF 9.050 34 2.2 FP 9.461 11 1.0 FP 4.988 35 4.5 FP 7.530 12 2.8 FP 5.643 36 2.9 FP 2.173 13 2.8 FP 7.348 37 3.9 FP 4.137 14 4.1 FP 4.160 38 .90 FP 6.357 15 1.0 FP 8.446 38 5.0 FP 4.798 16 1.2 FP 6.826 40 1.0 WF 7.819 17 2.1 FP 6.506 41 1.0 WF 8.184 18 3.6 FP 4.653 42 1.0 WF 5.063 19 4.4 FP 4.909 43 1.0 WF 5.131 20 0 FP 3.266 44 1.0 WF 2.507 21 4.8 SC 6.400 45 1.8 WF 6.053 22 4.8 SC 3.263 46 1.6 NF 6.169 23 4.5 SC 3.693 47 1.2 NF 6.105 24 4.6 SC 4.602 48 1.6 NF 3.949 49 .50 NF 5.385 O CO CO ^- ^ ( I I II I II — I I 1 1 ■H I Mil I SSMI Moisture Ordination Fig. 2. One-dimensional ordination by SSMI values of stands along the moisture gradient. 1979 Bloss. Brotherson: Moisture Gradient 167 life form groups and the moisture gradient was of interest, six groups were chosen— i.e., shrubs, succulents (cacti and cactuslike plants'), trees, woody plants, forbs, and se- lected plant families (i.e., composites and le- gumes). Figure 4 shows great similarity be- tween the established pattern for trees and that of legumes. It should be noted that the number of species these two groups had in common was high. Of further interest is the fact that the two groups contain species char- acteristic of the north-facing slope and of me- 30 ^ >• Opum, a fulgida -20 . 1 n 1 ■ Fercaclus wislizeni Fransena ambrospoides 0 1 2 « \ Fouguieria splendens Larrea divancata 20. 10. 0 Fransena delioidea ^40. 3-30. o20. <.o. Ce 40, 20. 1 2 ^^S Bacchans sarothroides 0 1 s o. SSMI Moisture Ordinal lon Polypogon monspeliensis 15 Hymenoclea mo nogyra 10. IS. 10. 5 . 0. SSMI VI oisiure Ordination S SMI Moist ure Ordination Fig. .3. Average percent frequency for indicator species plotted against the moisture gradient. 168 Great Basin Naturalist Vol. 39, No. 2 dium moisture regime preference. The shrubs, succulents, forbs, and woody plants as groups (Fig. 4) tend to prefer the dryer mois- ture habitats. The composites (as well as oth- er family groups such as the grasses) showed no preference for particular sections of the moisture gradient and exhibited, as groups, rather ubiquitous distribution patterns in relationship to the SSMI gradient. When cluster analysis techniques were ap- phed to the 37 species (Fig. 5), five major groups were apparent; however, a few spe- cies did not relate well to any group. From left to right on the dendrogram the groups are: (1) predominately flood plain and lower bajada species, which include Acacia greggii, Larrea divaricata, Schismus harhatus, Cerci- dium microphyllum, and Carnegiea gigantea; (2) north- and west-facing slope species, which include Fougiiieria splendens and Eriogonum fasciculatwn; (3) south- and east- facing slope species, which include Enchino- cereus Engehnannii and Opuntia leptocaulis; (4) flood plain species, which include Bromus rtibens and Franseria ambrosioides; and (5) the stream channel species Cynodon dacty- lon, Salix spp., and Hymenoclea monogyra. The relationships of these five groups to the established moisture gradient are depicted in Figure 6. As can be seen, all sections of the proposed moisture gradient are represented. When the dendrogram (Fig. 7) for the 49 stands was constructed, five groups were again recognized. Table 3 indicates how these groups relate with regard to the topog- raphy of the study area. From the cluster analysis it appears that the west-, south-, and east-facing slopes were highly similar in veg- etation types, and that those stands which were subjectively picked as being similar at the beginning of the study are in fact grouped together by cluster analysis. This 200 _ Shrubs 160 . 120 . 80 . 60 . 0 500 400. Woody Plants 300 . 200. 100. 60 40 . 20. 0. Composites h CO c 0) y Forbs 00 Trees 60. 40. 10. 0 . 0 3 4 5 ■ Legumes 120. 1 00. 80. 60. 20. 0. D 5 SSMI Moisture Ordination Fig. 4. Average percent frequency of life form groups plotted against the moisture gradient. June 1979 Bloss, Brotherson: Moisture Gradient 169 % Similarity — ^— Franseria deltoidea Cercidium microphyllum Plantago purshii Simtnodsia chinensis Carnegiea gigantea Opuntia bigelovii Opuntia engelmannii Lycium fremontii Acacia greggii Schismus barbatus Larrea divaricata Opuntia fulgida Prosopis juliflora Ephedra viridis Ferocactus wislizeni Eriogonum fasciculatum Fouquieria splendens Opuntia leptocaulis Eriogonum sp Mamnnillaria microcarpa Enchinocereus engelmani Salsola kali Lepidium sp Bromus rubens Franseria ambrosioides Baccharis sarothroides Cynodon dactylon Polypogon monspeliensis Salix sp Euphorbia sp Hymenoclea monogyra Datura meteloides Mimulus glabratus Baccharis glutinosa Platanus wrightii Marrubium vulgare Avena fatua >- Fig. 5. Dendrogram of "cluster" analysis of 37 species. 170 Great Basin Naturalist Vol. 39, No. 2 would, of course, be expected if one assumes that the vegetation of an area reflects the general pattern of its abiotic environment (Niering et al. 1963). Figure 8 illustrates niche width relation- ships to moisture preference. Flood plain spe- cies exhibit the broadest niche widths when compared with slope and stream channel spe- cies. Cluster analysis (Fig. 5) grouped species with similar niche widths together. Table 4 ranks all species in order of decreasing niche width and further serves to illustrate the point that those species with the broadest Table 3. Dendrogram groups as designated in Figure 7, along with included stands and their predominant as- pect and/or topographical location. % Similarity Dendrogram Predominant Group # Stand # location 1 21, 22, 23, 24, 25, 26, 28, 29, 30, 39 River channel 2 4, 11, 12, 13, 14, 18, 31, 32, 36, 37 Floodplain 3 1, 2, 3, 5, 6, 7, 8, Slopes (east. 9, 10, 15, 16, 17, 33, south, west) 40, 41, 42, 43, 44 4 20, 38, 19, 34, 35 Floodplain 5 45, 46, 47, 48, 49 North-facing slope Fig. 7. Dendrogram of "cluster" analysis of 49 study plots. Group 5 ^_^ Group 3 Group 1 Group 2 I I I Group 4 SSMI Moisture Ordination Fig. 6. Average frequency of cluster group species plotted against the inferred moisture gradient. 1979 Bloss, Brotmerson: Moisture Ci 171 niches are, generally speaking, flood plain species. As indicated earlier, the flood plain includes a wide variety of habitats and it may be this fact that influences the broadest nich- es being noted in the flood plain. Table 5 gives figures for the means (x) and standard deviations for NW, SSMI, and DI figures. Regres.sion analyses indicated that the NW vs. SSMI values exliibited highly signifi- cant correlations (p < .001). When niche 30, 25. £ 20 15 (V o Z 10 5. flood plain species slope species stream channel species Increasing moisture preference areas—* Average diversity index values and average niche width values plotted against the moisture gradient. 24 22 20 18 16 I - ^ 10 1000 2000 3000 Presence x Frequency Index 4000 5000 Fig. 9. Regression analysis of the relationship of NW vs. P X F index. 172 Great Basin Naturalist Vol. 39, No. 2 width measurements were compared by re- ters generally express similar information in gression analysis to the P X F index values terms of species importance in a community. (Table 4 and Fig. 9), the relationship was The relationship of the Diversity Index again highly significant (p < .001) with an r- (DI) parameter to stand placement along the value of .68. It appears that the two parame- moisture gradient is illustrated in Fig. 10. As Table 4. List of species encountered in the study, along with their niche width, percent presence, average per- cent frequency, and presence times frequency (P X F) index. Species are ordered in decreasing value of niche width. Niche Percent Average PxF Species width presence frequency index Acacia gre^ii Franseria deltoidea Cercidium microphyllum Plantago purshii Carnegiea gigantea Bromus rubens Schisinus barbatus Larrea divaricata Simmondsia chinensis Lyciiim fremontii Opuntia engelmannii Opuntia begelovii Hymenoclea monogijra Lepidiiim sp. Prosopis juliflora Baccharis sacrothroides Fotiquieria splendens Polijpogon monspeliensis Euphorbia sp. Cynodon dactylon Franseria ambrosioides Mammillaria microcarpa Ferocacttis wislizeni Datura meteloides Ephedra viridis Enchinoceretis engebnannii Opuntia fulgida Baccharis glutinosa Eriogonum fasciculatum Marrubium vulgare Opuntia leptocaulis Eriogonum sp. Salsola kali Mimuhis glabratus Avena fatua Salix sp. Platanus wrightii Table 5. Mean (X) and standard deviation (SD) values for the diversity indices (DI), synthetic stand moisture in- dices (SSMI), and niche widths (NW) as computed for stands grouped in cluster analysis (Figure 7 and Table 3). 22.359 73.5 67.4 4954 22.293 57.1 45.9 2620 19.724 42.9 55.2 2368 18.720 49.0 27.4 1343 16.529 38.3 41.7 1618 16.152 42.9 13.9 596 15.962 46.9 28.6 1341 15.949 36.7 55.2 2044 15.657 .30.6 70.6 2160 15.314 32.7 25.4 831 13.648 18.4 40.3 742 10.483 28.6 61.6 1742 8.389 22.4 48.1 1077 7.886 20.4 10.8 220 7.227 18.4 62.7 1154 7.184 14.3 42.4 606 5.450 8.2 57.8 66 5.391 18.4 18.5 340 5.181 14.3 17.8 225 4.518 18.4 22.2 408 4.466 16.3 19.6 319 4.000 8.2 57.8 474 4.000 14.3 20.1 287 4.000 6.1 8.3 51 3.006 6.1 8.3 51 2.986 12.2 15.6 190 2.910 20.4 46.3 945 2.575 6.1 66.5 404 2.571 8.2 31.2 256 2.000 4.1 8.3 34 1.853 6.1 27.7 169 1.579 4.1 16.7 68 1.577 4.1 16.7 68 1.293 16.3 9.4 153 1.000 2.0 8.3 17 1.000 2.0 33.0 66 1.000 2.0 33.0 66 DI SSMI NW Groups' X SD X SD X SD 1 3.67 1.27 4.25 1.18 8.36 5.82 2 4.58 1.74 2.98 1.46 13.90 6.84 3 6.47 1.73 1.02 .31 14.98 6.44 4 6.30 2.35 2.40 2.03 12.48 6.32 5 5.54 .97 1.34 .52 11.69 7.41 June 1979 Bloss, Bhotherson: Moisture Gradient 173 opposed to niche widths, stand diversity was highest in the bajada and slope stands. In tropical ecosystems high uniformity is posi- tively correlated with high diversity (Mac- Arthur and Wilson 1967). However, in north temperate systems this relationship appears to be reversed— i.e., low uniformity yields high diversity (Dick 1971, Hall 1971). The data from our study seem to follow a similar pattern. There are, however, other parame- ters not measured in this study (i.e., disturb- ance as indicated by the presence of the in- troduced species Avena fatua and Brotniis rubens, etc.) which, it is felt, have influenced the diversity trends of the area. Conclusions It is apparent that the stream channel has a set of species (among which are Polypogon monspeliensis. Datura meteloides, Baccharis glutinosa) distinct in many ways from those in the other areas; yet all are clearly a part of an integrated system. For example, Bromus rubens, Cercidium microphyUum, and Acacia greggii grow not only in the stream channel 35 30, 25. 10. 5. 0 12 3 4 5 SSMI Moisture Ordination Fig. 10. Niche width (NW) plotted against species ar- ranged by dendrogram sequence. but also on the flood plain and slopes. The flood plain is highly varied in habitat, result- ing in a mosaic of types. Some of the stream channel vegetation intergrades only into the flood plain, and the flood plain vegetation in turn intergrades into slope vegetation, espe- cially where the plain is wide. In some loca- tions where it is narrow the change between flood plain and channel vegetation or be- tween flood plain and slope plain is sharply delineated, so that one can easily delineate channel food plain or slope vegetation. In other areas the flood plain is broad and the change from channel to flood plain to slope is so gradual and smooth that one is hard put to find where the stream channel ends and the flood plain begins. Due to the wide variety in habitats on the flood plain those species that are found to be ubiquitous throughout the flood plain region exhibit the broadest niche. Such species (i.e., Acacia greggii, Cercidium microphyUum, Larrea divaricata) with broad niche widths were predominately species of importance in the flood plain and lower ba- jada. Where the slope is distinct from the flood plain, it is clearly more sparsely vegeta- ted than the flood plain and is dominated by several small shrubs. Franseria deltoidea is most prominent in these areas. In still other locations there exists a mixture of many spe- cies wherein no one species distinctly domi- nated. Species of similar life forms and taxa often showed similar location preference. Family groups in general, however, were ubiquitous and everywhere present. For example, the composites, as a family, are probably the most prevalent group in the study area and showed no identifiable trends. Other promi- nent families are the Poaceae and the Fa- baceae. The grasses, though ubiquitous in dis- tribution, showed no decided location preference. The legumes, on the other hand, appear to prefer medium moisture areas. With many of the legumes being trees or treelike forms, the arbrorescent life form also exhibited medium moisture preferences. Sur- prisingly, the forbs preferred the dryer mois- ture areas; however, this may be due to fac- tors other than moisture. Succulents which are predominately cacti preferred the dryer moisture regimes. Carnegiea gigantea, one of the more prominent cacti on the area, exhib- 174 Great Basin Naturalist Vol. 39, No. 2 ited its highest frequency value on the slopes. It did, however, extend onto the flood plain in some locations. Shmbs, as a group, repre- sented by species like Franseria deltoidea, Larrea divaricata, and Franseria amhrosioides also preferred the dryest moisture areas. Cluster analysis results were useful in that they substantiated many assumptions of the initial hypothesis which were subjectively stated. Species when clustered together ex- hibited similar niche widths and were also observed to occupy habitats in similar topo- graphical locations. Stands when clustered produced groups which correlated to all sec- tions of the moisture gradient. This evi- denced that the gradient (though synthet- ically constructed) did, in fact, reflect natural conditions. Six different areas were estab- lished and delineated as distinct (i.e., stream channel, flood plain, north-, west-, south-, and east-facing slopes). Cluster analysis in- dicated distinctness in the flood plain, stream channel, and north-facing slope stands, but clustering west-, south-, and east-facing slope stands were as one imit. Vegetatively speak- ing, these three areas were highly similar and support vegetation distinctly different from the other three locations. The patterns exhibited when indicator spe- cies frequency values were plotted against the moisture gradient also substantiate that the established moisture gradient is realistic and that the indicator species chosen are good index plants of general moistvire condi- tions. The relationships noted between the P X F index and the niche width measurements indicate that the two express similar informa- tion in terms of species importance in a sys- tem. Their correlation to each other is posi- tive and highly significant. Diversity was shown to be highest on the slopes and negatively correlated to moisture (i.e., as moisture increases diversity decreases) (Fig. 11). It is felt, however, that the diversity data would have been more meaningfully in- 3. slope vegetation stream channel vegetation Mean SSMI Fig. 11. Regression analysis of relationship between DI and SSMI. June 1979 Bloss, Brothkhson: MoisTuuE Gradient 175 terpreted if other environmental parameters (i.e., disturbance, etc.) had been measined. Regression analysis of DI vs. SSMI exliibited a negative correlation at .001 level of signifi- cance and had an r- value of .74. Literature Cited Anderson, O. 1964. The phytosociology of dry lime prairies of Wisconsin. Unpublished dissertation, University of Wisconsin. Barbour, M. G. 1968. Germination requirements of the desert shrub Larrea divaricata. Ecology 49:915-923. Beatley, J. C. 1967. Survival of winter annuals in the northern Mojave Desert. Ecology 48:745-750. Blumer, J. C. 1910. A comparison between two moun- tain sides. Science 31:834-840. 1909. On the plant geography of the Chiricahua Mountains. Science 30:720-724. Brians, K. 1928. Change in plant association by change in ground water level. Ecology 9:474-478. Capon, B., and W. V. Asdell. 1967. Heat pre-treatment as a means of increasing germination of desert annual seeds. 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Relation of soil moisture to desert vegetation. Bot. Gaz. 50:241-256. MacArthur, R. H. 1972. Geographical ecology; patterns in the distribution of species. Harper and Row, New York. 251 p. MacArthur, R. H., and E. O. Wilson. 1967. The theo- ry of island biogeography. Princeton University Press, Princeton, New Jersey. 199 p. Mallery, T. D. 19.36a. Rainfall records for the Sonoran Desert. Ecology 17:110-121. 1936b. Rainfall records for the Sonoran Desert. II. Summary of readings to December 1935. Ecol- ogy 17:212-215. Marks, J. B. 1950. Vegetation and soil relations in the Lower Colorado Desert, eco. 31:176-193. MuLLER, W. H., AND C. H. MuLLER. 1956. Associatiou patterns involving desert plants that contain tox- ic products. Amer. J. Bot. 43:354-360. NicHOL, A. A. 1952. The natural vegetation of Arizona. Univ. of Arizona Agric. Expt. Sta. Tech. Bull. 127:189-2,30. NiERiNG, W. A., R. H. Whittaker, C. H. Lowe. 1963. The Saguaro: a population in relation to environ- ment. 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Physical conditions of sun and shade. Ecol- ogy 12:96-104. Smith, H. V. 1956. The climate of Arizona. Univ. of Ari- zona Agric. Expt. Sta. Bull. 279:1-99. Sneath, p. H. a., and R. R. Sokal. 1973. Numerical tax- onomy. W. H. Freeman and Co., San Francisco. 573 p. SoRENSON, T. 1948. A method of establishing groups of equal aptitude in plant sociology based on sim- ilarity of species cont. Konge. Dan. Vidensk. Selsk. J. S. 4:1-34. Spalding, V. M. 1909. Distribution and movement of desert plants. Carnegie Inst. Wash. Publ. 113. 114 p. 1910. Plant associations of the Desert Laboratory Domain and adjacent valley. Plant World 13:31-42. Trevis, L. J. 1958a. Germination and growth of ephem- erals induced by sprinkling a sandy desert. Ecolo- gy 39:681-688. 1958b. A population of desert ephemerals germi- nated by less than one inch of rain. Ecology 39:689-695. Turner, R. M. 1963. Growth in four species of Sonoran Desert trees. Eco. 44:60-65. Turner, R. M., S. M. Alcorn, G. Olin, and J. A. Booth. 1966. The influence of shade, soil and water on saguaro seedling establishment. Bot. Gaz. 127:95-102. Walmo, O. C. 1955. Vegetation of the Huachuca Moun- tains, Arizona. Amer. Midland Natur. 54:466-480. Went, F. W. 1942. The dependence of certain annual plants on shnibs in southern California deserts. Bull. Torrey Bot. Club 69:100-114. 1948. Ecology of desert plants. 1. Observations on germination in Joshua Tree Nat'l Mon., Gal. Ecology 29:242-253. 1949. Ecology of desert plants. II. The effect of rain and temperature on germination and growth. Eco. 310:1-13. Went, F. W., and M. Westergaard. 1949. Ecology of desert plants. III. Development of plants in Death Valley Nat'l Mon., Gal. Ecology 30:26-38. Whittaker, R. H., and W. A. Niering. 1964. Vegeta- tion of the Santa Catalina Mountains, Arizona. I. Ecological classification and distribution of Spe- cies. Arizona Acad. Sci. J. 3:9-34. 1965. Vegetation of the Santa Catalina Moun- tains, Arizona: A gradient analysis of the south slope. Ecology 46:429-452. WiLM, H. G. 1956. Treatment of vegetation on the Salt River watershed to increase water yield. In: Re- covering rainfall. Ari. Watershed Comm. p. 209-218. Yang, T. W. 1950. Distribution of Larrea tridentata in the Tucson area as determined by certain phys- ical and chemical factors of the habitat. Unpub- lished thesis. University of Arizona. 1957. Vegetational, edaphic and faunal correla- tions on the western slope of the Tucson Moun- tains and the adjoining Avra Valley. Unpublished dissertation. University of Arizona. Yang, T. W., and C. H. Lowe. Correlation of major veg- etation climaxes with soil character in the Sono- ran Desert. Science 123:542. ECOLOGICAL AND COMMUNITY RELATIONSHIPS OF ERIOGONUM CORYMBOSUM (POLYGONACEAE) IN THE UINTA BASIN, UTAH Jack D. Brotherson' and Karen J. Brotherson' Abstract.— Ecological and community relationships of 10 different plant communities in the Uinta Basin, Utah, where Eriogonum corymbosum was found to grow were studied and described. Each commimity was sampled to determine the amount of ground cover, percent composition, frequency, and density of participating species. Phys- ical site factors, viz., soil texture, total soluble salts, pH, cation exchange capacity, and amounts of calciinn, magne- sium, potassium, and sodium were determined. The 10 communities were compared to determine the degree of sim- ilarity between them. Correlations between individual plant species and measurable characteristics of the community were attempted. Evidence is presented that the distributional patterns of some species are related to these measured characteristics. Eriogonum corymbosum, Chenopodium leptophyllum, Atriplex con ferti folia, Stipa comata, Artemisia tridentata, and Agropyron smithii showed correlation to both vegetational and edaphic factors of the community. Total vegetative cover increased from desert to mountain in the Uinta Basin. As the vegetative cover increased, soil depth also increased. Eriogonum corymbosum decreased in importance in the higher elevation communities. Eriogonum corymbosum was studied taxonomically, which demonstrated the presence of a previously undescribed variety. It is suggested that E. corymbosum var. corymbosum, found generally in the desert areas of the basin, is composed of a series of ecotypes that inhabit shallow soils and prefer communities that show high degrees of disturb- ance, little competition, fairly high levels of soluble salts in the soil, and are found at elevations below 5500 feet. Eriogonum corymbosum var. erectum, on the other hand, does best in communities above 6000 feet that show less disturbance than the desert areas, have deeper soils, and low levels of soluble salts. Eriogonum corymhosiim Benth. in DC (wild buckwheat) is a low-growing, perennial shmb that occurs in much of the cold tem- perate desert shrub regions of Colorado, Utah, and Nevada. In the Uinta Basin its growth is widespread on several different geological formations. In some areas its dis- tribution is restricted to a specific formation, while elsewhere its distribution appears un- restricted. The ecology of E. corymbosum is not well known, and available literature on the subject is fragmentary and scattered. Welsh (1957), in a study of the Dinosaur National Mon- ument in the Uinta Basin, Utah, describes its occurrence on the Moenkopi and Mowry shale formations. He states that it appears to be restricted to these formations and grows only where the formations are fully exposed. It is the dominant plant of much of the Mow- ry Shale formation. In an ecological study of the Flaming Gorge Reservoir Basin, Flowers (1957) lists E. corymbosum as a member of a zone of vege- tation forming a junction between the river banks and the uplands, where it appears fre- quently on dry hillsides at about 1700 m (5600 feet). Graham (1937) found it growing with sagebrush on Red Creek. Many writers have noted that plants are indicators of certain soils and geological for- mations. Graham (1937) wrote of the exist- ence of endemics on the Green River shale formation in the Uinta Basin. Cannon (1952) noted the correlation between uranium-van- adium deposits and vegetation. Mason (1946) states that some soils, such as the serpentine soils, are well known for the endemic species that occur on them. Krucheberg (1951, 1954), Whittaker (1954), and Waler (1954) note a correlation between endemics and indicator species of the serpentine soils of California. Kearney et al. (1914) in Tooele Valley, Utah, found that different types of native vegetation indicated conditions of moisture and salinity of the soil on which they were 'Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 177 17J Great Basin Naturalist Vol. 39, No. 2 found. Shantz and Piemeisel (1940) found conditions in Escalante Valley, Utah, similar to those found in Tooele Valley, Utah, by Kearney et al. (1914). Fautin (1946) noted shadscale scattered in the more xeric and higher saline areas of valleys in central Utah. Billings (1949, 1950, 1951, 1952) states that chemical differences in soils may produce marked differences in the vegetation within fairly uniform climatic areas, that vegetation zones can be correlated with climatic and soils factors, and that the mosaic of smaller vegetational differences within a large zone may be caused by edaphic factors. He also noted Atriplex confertifolia to be an indicator of subsoil salinity. Gates et al. (1956) consid- ered the edaphic factors, soluble salts, satu- rated extract conductivity, and exchangable sodium, to be of primary importance in de- termining the distribution of some shrub types. Thatcher (1959) and Robertson et al. (1966) found that Atiemisia tridentata oc- curred only on moderately deep to deep, loamy soils. Little has been published on the subject of indicator species concerned with the cold temperate desert shrub regions of the west. A knowledge of indicator species such as E. co- rymbosum is desirable because this species grows on peculiar soil types (i.e., soils of high or low pH, high or low salt content, etc.) and thus becomes important to the ecologist at- tempting to delineate or describe areas of the cold temperate regions. During this study, I recognized from an ecological standpoint two distinct varieties. Consequently, the two varieties were named E. conjmbosum Benth. in DC var. conjmho- sum and E. conjmbosum Benth. in DC var. erectiim Reveal and Brotherson (Reveal 1967). Eriogoniim corymboswn var. conjmbosum has its distribution centered mainly in north- eastern and central Utah, but it extends into northwestern Colorado and extreme south- western Wyoming. It also extends southward to northcentral and northwestern Arizona. Eriogonum conjmbosum var. erectum is widely scattered in northeastern Utah, ex- tending from western Wasatch County to ex- treme western Uintah County, mainly above 1800 m (6000 feet) elevation. Study Area The Uinta Basin is a broad, elongated, asymmetric basin lying in the northeastern corner of Utah and extending into north- western Colorado. It is both a stRictural and a topographic basin and is a subdivision of the Colorado Plateau province (Marsell 1964). It encompasses some 12,000 square miles and is about 150 miles (240 km) long in an east-west direction and 100 miles (160 km) wide in a north-south direction (Dastrup 1963). The basin's northern boundary (Fig. 1) is in the Uinta Mountains, which reach elevations of 4100 m (13,500 feet). On the south it is bounded by the rim of the Tavaputs Plateau, which appears to rise roughly but steadily to a dip slope. Its west rim is flanked by the eastern slopes of the Wasatch Mountains, but its eastern side is not so sharply defined topo- graphically (Marsell 1964). Much of the central portion of the Uinta Basin is desert with an annual precipitation of less than 24 cm (10 inches). The gently sloping floor lies between 1500 m (5000 feet) and 1800 m (6000 feet) in elevation, but it reaches as much as 2100 m (7000 feet) at the foot of the Uinta Mountains. The basin is drained in a somewhat centri- petal pattern by three major streams and their tributaries (Fig. 1). The Duchesne River flows in a southeast direction and the White River flows westward. These two streams, along with their tributaries, drain the entire basin excepting the northeast corner. This corner is drained by the third major stream, the Green River, which crosses the East Tavaputs Plateau to meet the Colorado. The Duchesne and its tributaries (Rock Creek, Yellowstone River, Lake Fork, and the Uinta River) drain almost the entire Utah part of the basin, and the White River drains the Colorado portion. The Strawberry River, found in the western end, is the only stream of any size that drains the southern part of the basin (Marsell 1964, Dastrup 1963). The land surface of the basin is rough and broken, cut in many places by deep gorges. Bare rock surfaces are widely scattered, forming cliffs, deep slopes, and stripped sur- faces in many places. The soil is highly vari- able, ranging from deep, heavy clays to shal- June 1979 Brotheuson, Bhothehson: Plant Community 179 low, sandy soils. In much of the basin a calcareous layer can be found (Marsell 1964). Geology The Uinta Basin is a typical Rocky Moun- tain-type assymetric Tertiary basin. It was formed by the gradual .settling of the interior area below the surface of deposition and con- current lesser sinking or upward movement of the basin rims. Its development began in the Paleocene or Eocene and has continued until the present. As before stated, the basin is both a topographic and a structural basin. It exhibits from 900 m (3000 feet) to 1800 m (6000 feet) of relief between its lowest basin parts and the highest rim. The basin repre- sents a superficial expression of the Tertiary structural basin and would still be filling with sediment except for the Green River, which has breached the north and south rims (Os- mond 1964). The basin contains a large number of geo- logical formations, each having its own age and period of development. Seven of these formations furnish substrata for the 10 differ- ent communities analyzed in this study. These 7 formations (in sequence from the oldest to the youngest) are: Ankareh, Moen- kopi, Mowry Shale, Parachute Creek and Evacuation Creek (members of the Green River formation), Uinta, Duchesne River, and glacial moraines. Methods Ten study areas were established through- out the Uinta Basin (Fig. 1). The areas were chosen on the basis that Eriogonum conjmho- suin was present in the vegetational cover and by the geological formations upon which they occurred. Vegetational data and soil samples were collected during the summer MAP OF THE UINTA BASIN • TOWN LIMIT OF BASII Fig. 1. Map of the Uinta Basin showing study site locations. 180 Great Basin Naturalist Vol. 39, No. 2 months (June through August) and analyzed in the laboratory during winter. Vegetation data were gathered in midsum- mer (July 15 to August 10) when E. conjmho- sum and most other species had reached their maximum growth. Collecting of voucher specimens was done throughout the growing season. All specimens were deposited in the Brigham Young University herbarium. The vegetational cover of each area was analyzed by employing the line-point method (Cain and Castro 1959). Ten 15-m (50-feet) transects were taken in each area, with the points distributed every 7 cm (3 inches) along the transect. This gave a total of 200 points per transect or 2000 points per area. Trans- ects were located on a restricted random basis to eliminate bias and keep adjacent transects equal distances apart. At each point along the transect a seven-gage sharpened wire was lowered toward the ground and re- cords were made of the first and all succes- sive species hits. Where no vegetation oc- curred for the wire to touch, the hits were recorded as litter, rock, or bare ground. Frequency and density data were acquired by use of a meter square quadrat placed at 3- m (10-foot) intervals along the 15-m trans- ects. This gave a total of 50 quadrats per area. Frequency and density data were re- corded only for shrubs, trees, and bunch grasses. Percent surface cover for individual spe- cies and percent total vegetation cover were calculated as follows: surface cover Total hits for each species for each species Total points Total hits for all % total vegetational species minus overlapping hits cover Total points Total hits for % composition each species X 100 Percent frequency figures and density fig- ures were obtained through use of the follow- ing relationships: Total quadrats a species appears in % frequency Density /acre- X 100 Total quadrats # plants/50 quadrats X 83.00 50 quadrats Differences and similarities between the different study sites were computed utilizing Sorensen's index of similarity (Sorensen 1948), as adapted by Dix and Butler (1960). The formula was as follows: K = 2W/a -H b X 100. K is the similarity index between two sites; a represents the sum of the coverage percent- ages of all species in one stand; b represents a similar figure for a second stand. W repre- sents that part of the cover common to the species found in both stands. After the indices were obtained, an ordination matrix was con- structed (Table 4). This placed the most sim- ilar sites next to each other and the more dis- similar sites at opposite ends of the matrix. The sites were then divided into four groups and a table (Table 6) was constRicted from Table 1. Site characteristics areas. .'iated with 10 study of each species Total hits for all ■ species % Soil Exposed % % depth bare Exposed Litter Site (dm) ground rock cover Mowrv 1.86 57.80 0.00 2.15 Moenkopi 4.36 76.50 2.00 0.10 Evacuation Creek .98 59.00 9.40 4.10 Parachute Creek 1.77 56.50 6.25 2.75 Wells Draw 1.21 62.80 2.00 2.15 Red Creek 5.06 45.30 13.50 6.00 Rock Creek 3.32 13.80 7.80 6.60 Strawberry Valley 2.81 29.25 2.25 4.10 Uinta River 4.96 9.55 0.15 5.70 Yellowstone River 7.31 .90 0.00 4.25 'This number (83.00 50 quadrats/acre) was obtained by dividing the number of square feet in 50 quadrats (i.e., 524. f number of square feet in an acre (i.e., 4,3,360 square feet/acre). i square feet/50 quadrats) into the 1979 Bhothehson, Bhoiherson: Plant Community 181 Table 2. Soil data for each study site by horizon. Each figure represents an average for three pits at each site. Soluble CECin Depth Sand Silt Clay pH salts Meq./ Ca Mg K Na (in inches) % % % 1:1 (ppm) /lOOg. % % % % Mowry Format ion Surf. 65 17 18 6.4 1099 36.88 53.4 28.3 9.7 8.1 0-6 63 18 18 4.8 1295 46.59 57.0 28.6 6.4 8.0 6-12 64 13 14 3.8 1127 48.61 61.8 23.4 5.7 9.0 12-18 63 13 15 3.7 1023 44.66 62.8 22.3 5.7 9.5 Moenkopi Formation Surf. 41 — — 7.9 818 8.57 80.4 7.4 2.8 9.5 0-6 54 _ _ 8.0 887 8.83 79.8 7.0 4.1 9.0 6-12 62 — — 7.9 1214 9.29 72.1 9.2 6.1 9.9 12-24 65 _ _ 7.9 1440 9.54 70.0 10.3 7.2 10.5 24-36 61 - - 8.1 1875 8.84 68.2 13.0 7.5 10.8 Evacuation Creek Member Surf. 83 10 7 8.5 208 11. .39 69.4 12.1 4.2 14.4 0-6 72 18 10 8.7 256 16.20 70.8 12.5 4.1 12.6 6-12 74 11 16 8.8 267 16.35 65.5 14.3 3.3 17.0 Parachute Creek Member Surf. 71 22 8 8.3 530 20..33 66.7 21.1 2.8 9.1 0-6 58 26 16 8.3 550 24.03 58.6 29.7 2.3 9.1 6-18 68 25 7 8.1 630 25..38 55.6 34.4 1.9 8.3 18-24 — — — 8.2 373 Wells Draw — — — — — Surf. 78 18 3 8.4 196 13.27 76.9 7.9 6.9 8.2 0-6 69 23 8 8.5 234 16.75 76.1 10.5 5.5 9.8 6-12 62 23 10 8.2 401 2.3.71 70.9 12.5 3.3 13.4 12-24 80 16 4 8.5 229 Red Creek 17.94 78.1 9.9 1.6 10.4 Surf. 42 37 24 8.8 386 14.88 48.4 41.2 5.1 5.4 0-6 32 46 22 9.0 410 16.75 .38.1 50.9 4.3 6.8 6-12 27 47 26 9.2 461 16.62 31.8 54.5 5.2 8.5 12-24 32 43 26 9.5 519 14.75 .30.3 50.5 3.7 15.5 24-36 44 38 18 9.6 630 Rock Creek 14.08 28.1 49.8 4.3 17.8 Surf. 53 35 12 8.3 271 11.69 65.2 18.5 10.3 6.0 0-6 55 34 11 8.3 261 13.36 59.6 23.5 10.4 6.5 6-12 54 36 10 8.5 248 12.31 59.0 28.2 7.0 5.4 12-24 48 37 12 8.4 274 9.96 53.7 30.3 9.5 6.5 24-36 45 - - 8.2 924 5.38 61.9 31.0 7.9 7.4 Strawberry Vallev Surf. 51 29 20 8.2 305 8.83 62.7 20.4 10.0 6.9 0-6 50 30 20 8.3 357 8.63 58.5 26.9 6.9 7.9 6-12 43 38 19 8.5 279 10.63 55.6 32.9 4.3 7.1 12-24 51 34 15 8.7 211 Uinta River 6..35 55.3 34.3 4.3 6.1 Surf. 59 28 13 7.3 308 15.88 56.3 24.1 12.7 6.9 0-6 58 27 14 7.8 391 19..36 56.6 22.3 13.7 7.4 6-12 58 25 17 8.0 400 19.45 54.8 23.8 14.3 7.1 12-24 60 23 17 8.1 394 18.07 51.9 27.1 13.9 7.0 24-36 60 22 18 8.2 410 17.43 49.6 29.4 13.9 6.7 Yellowstone River Surf. 80 16 5 5.9 42 8.54 66.3 15.6 9.1 9.1 0-6 83 12 4 6.7 47 5.83 66.0 18.4 6.8 9.0 6-12 89 7 4 6.7 46 4.45 62.9 21.1 7.6 8.8 12-24 85 10 4 6.8 35 4.77 63.6 22.0 5.0 9.5 24-36 87 8 4 6.8 36 4.42 60.9 23.6 6.1 9.4 182 Great Basin Naturalist Vol. 39, No. 2 Piv these by averaging the cover percents for each species within the group. Those plant species showing the highest preference for group 1 were placed at the top of the table, and those species showing the least prefer- ence were placed at the bottom. From this table, 15 indicator species were selected and given adaptation numbers. Those species with the greatest preference for Group 1 were given the number one and those with the greatest preferences for Group 4 were as- signed the number five. By use of the selected indicator plants. Plot Index Values (PIV) for the 10 sites were ob- tained by applying the following formula (Dix and Butler 1960): Sum (relative composition of each indicator species X its adaptation #) Sum (relative composition of each indicator species) From the Plot Index Values a linear ordi- nation (Fig. 2) was constructed. Environmen- tal correlations were then attempted with the use of this ordination. A list was made of all plants noted in each area, and a composition study was made (Table 10) to indicate which families were the best represented in each area. All plant specimens collected during this study were deposited in the herbarium of the Brigham Young University. Three soil pits were dug at each study area, and soil samples containing a composite sample of several hundred grams were taken from the surface, 0-15, 15-30, 30-60, 60-90 cm depths. Soil depth was determined for each area by using pits and a 10-dm penetrometer (a sharpened %" steel rod). The penetrometer was pvished into the ground at 10-foot inter- vals along the 50-foot transects used to col- lect the vegetational data. Textural analysis of the soil was deter- mined according to Bouyoucos (1936, 1951). Hydrogen ion concentration was determined for the same group of samples on a saturated paste, a 1:1 and a 1:5 ratio of soil to water. The 1:1 and 1:5 ratios were measured to in- dicate sodium content. A Beckmann glass electrode pH meter was used and the samples were prepared as outlined by Russell (1948). Total soluble salts for the above-mentioned saturated pastes were determined by the use of a Wheatstone electrical conductivity bridge. Cation exchange capacities for all collect- ed soil samples were determined by using the standard ammonium acetate extraction meth- od described by Russell (1948). Analysis for the amount of available calcium, magnesium, potassium, and sodium was found by using the ammonium acetate extraction from the cation exchange capacity determinations and a Beckmann model DU flame photometer. Results and Discussion The 10 study sites were located throughout! the Uinta Basin (Fig. 1). General physical andl biological factors characteristic of these sites* can be seen in Tables 1, 2, 3, and 8. Those sites located in the south and eastern parts of the basin showed shallow soils, little litter cover, and large exposed areas of bareL ground. In contrast, those sites located in thef western and northern parts of the basin ex- hibit deeper soils, increased amounts of littei cover, and much less exposed soil surface (Table I). Texturally the soils varied from sand tc loams to sandy clay loams, with no distinct patterns being evident (Table 2). Soluble sail concentrations and cation exchange capaci ties generally increased from the north anc 1 3' 24 5 J L I I Plot Index Ordination Fig. 2. The plot index values ordination. The linear arrangement of the 10 study sites according to their plot iiide> values. 1979 Brotherson, Brotmerson: Plant Community 183 west parts of the basin toward the south and east parts. Hydrogen ion concentration (pH) varied from lows of 3.7 on the Mowry Shale formation to 9.6 in Red Creek on the Uinta formation, but again showed no distinct geo- graphical patterns. Calcium and magnesium were the most abundant of the cations mea- sured, with potassium and sodium seldom comprising over 10 to 20 percent of cation exchange capacity (Table 2). Vegetationally the sites located in the north and west parts of the basin showed the highest shrub densities, even though grasses were the dominant life form type with regard to cover. Shrub cover, on the other hand, was the most important in the south and east I parts of the basin (Table 3). Forb and annual I plant cover showed no observable patterns. Total cover and diversity patterns (Table 3), in contrast to bare ground (Table 1), were highest near the mountains in the north and west basin areas. Numbers of species per study site varied from 11 to 29, with an aver- age of 22. The prevalent species (the 24 most prominent species found to occur across the study area) are listed in Table 8. It can be seen they represent a rather typical assem- blage of the cold temperate desert shrub areas of the intermountain region. Table 5. Study site groups as determined from the similar index matrix. Group Study site Mowry study site Parachute Creek member of the Green Fliver Formation study site Evacuation Creek member of the Green River Formation study site Moenkopi study site Red Creek study site Rock Creek study site Strawberry Valley study site Uinta River study site Yellowstone River study site Table 3. Vegetation characteristics associated with the 10 study sites. Site Total Percent cover Shrub density plants/Ac cover Shrub Forb Grass Annual 40.1 78.0 17.0 4.0 1.0 7300 20.5 81.0 15.0 3.0 1.0 2515 27.5 77.0 4.0 7.0 12.0 8449 34.5 64.0 1.0 32.0 3.0 4542 33.5 68.0 1.0 20.0 11.0 7138 34.8 68.0 0.5 .30.0 2.0 15572 72.6 28.0 3.0 65.0 4.0 15660 64.4 39.0 13.0 45.0 3.0 24652 84.6 37.0 9.0 51.0 4.0 37716 94.9 22.0 4.0 60.0 14.0 7301 Mowry Moenkopi Evacuation Creek Parachute Creek Wells Draw Red Creek Rock Creek Strawberry Valley Uintah River Yellowstone River .53 "Diversity = 1/2 pi' Table 4. Similarity index matrix for the 10 study sites: Mowry = 1; Moenkopi = 2; Evacuation Creek = 3; Parachute Creek = 4; Wells Draw = 5; Red Creek = 6; Rock Creek = 7; Strawberry = 8; Uinta = 9; Yellowstone = 10. 10 48.0 45.6 45.0 40.6 32.1 20.8 17.1 17.5 11.7 57.6 50.5 46.0 ,35.2 25.3 21.2 17.3 14.7 62.4 45.7 .35.5 22.5 18.9 17.8 12.9 50.2 .39.1 2,3.1 17.6 15.7 11.6 31.7 29.8 25.6 18.3 13.2 50.6 21.6 24.2 16.2 27.8 25.5 17.5 29.3 33.5 62.2 1S4 Great Basin XatuR-alist Vol. 39. No. Ordination .\nalysis -\nalvsis of the vegetation of the different areas and of the areas themselves was at- tempted bv evaluating the differences and similarities between sites. To accomplish this, the areas were ordered by application of an index of similarit\" Table 4 . The matrix was designed to place those sites with the greatest similarities closest to- gether and those with the greatest differences farthest apart. Therefore, the Moenkopi and the Evacuation Creek sites are interpreted to be the most similar and will thus exhibit sim- ilar communitv characteristics, and the Moenkopi and the Yellowstone sites are the most dissimilar and should exhibit few factors in common. The position of each area within the matrix was detennined by placing the highest index values along the diagonal of the square and the lowest index values toward the upper right-hand comer of the table. The magnitude of the indices decreases from left to ri^ht when comparing the Mowr\' stud\" site with the Yellowstone River study site or the other sites listed to the ri2;ht of the Mow- r\". Similarly, the indices increase from top to bottom or from right to left when the other areas are compared to the Yellowstone River studv site. Table 6. Average percent composition values in the four groups for all species with a percent composition greater than 0.50 percent. Plant species Groups Atriplex confertifolia Chenopodiuum leptophyllimi Gila leptomeria Ephedra torreyana .Astragalus saurinus Juniperus utahensis Stanleya pinnata Ej-iogoniun cor\Tnbosum Tetrad>-mia spinosa MentzeUa dispersa -\melanchier utahensis Poa secimda Chpt^sothamnus panyi C)T\:zopsis h\Tnenoides Agrop\Ton trachycaulum Chrvsothamnus \Tscidiflorus .\rtemisia nova Lepidium montanum Sohdago petradoria Poa palustris Tetrad\Tnia glabrata .\stragalus tenellus Cercocarpus montanus Sitanion h\-stris .\rtemisia tridentata Purshia tridentata .Astragalus convallarius Sphaeralcea coccinea Lappula redowsldi Chr\3othamnus nauseosus Opuntia polyacantha Lepidium densiflorum Xanthocephalum sarothrae -\grop\Ton smithii Stipa comata 5.6S 4.95 3.42 2.01 61.05 0.25 1.28 3.91 0.27 0.12 1.78 4.3: 2.90 1.10 3.07 2.20 1.73 48.78 4.96 0.68 1.28 0.37 7.01 0.24 1.50 9.60 1.31 0.87 0.18 0.97 0.43 M.40 1.10 0.3S 4.94 33.10 1.03 14.16 0.16 6.35 0.48 0.46 O.OS 018 0.07 1.66 8.51 0.25 1.40! 0.S6 7.98 0.16 5.70 1.2c 1.3f 3.15 0.3t' S.31 0.2.e l.OC 1.45 0.31' 1.3- 1.3- i.o: 4.2^ 20.6- 26.1; These numbers are adaptatioa numbers and were assigned to different preference for a certain gnx^ or groups as diown in the above table. indicatOT species. Tbe indicatof species were cbosen on the baas of tfaeir indjcatw June 1979 Brotjierson, Brotherson: Plant Community 185 To help understand the association of indi- vidual species to these relationships, the areas listed on Table 4 were divided into the four groups shown in Table 5. The percent composition figures of all spe- cies with percents greater than 0.50 percent were then averaged in each group. A list (Table 6) was then prepared placing those species with the greatest preference for Group 1 at the top and those species with the greatest preference for Group 4 at the bot- tom of the list. You can now determine some Table 7. Plot index values (PIV) for 10 study sites. Study Site PIV 1 Mowrv 4 Parachute Creek 3 Evacuation Creek 2 Moenkopi 5 Wells Draw 6 Red Creek 7 Rock Creek 8 Strawberry Valley 9 Uinta River 10 Yellowstone River 1.86 2.08 1.95 2.03 2.21 3.14 3.28 3.65 4.51 4.37 characteristic distribution or associational patterns for many of the individual species. Although Table 4 served effectively to seg- regate the different study areas, it did not show the degree of compositional differences or similarities actually existent between them. Because such information was desir- able. Plot Index Values (PIV) (Table 7) de- scribed by Dix and Butler (1960) were em- ployed to assign a spatial position to each study site on a linear ordination and thus to actually measure to some degree the ecologi- cal distance between the different areas of study. To accomplish this, 15 indicator spe- cies were chosen from the list in Table 6 and assigned adaptation numbers from 1 to 5. Those species with the greatest preference for Group 1 were assigned the adaptation number 1, and those with the least prefer- ence for Group 1 were assigned adaptation number 5. Under such a system, an area will have a PIV of 1 when it contains only in- dicator species assigned adaptation numbers of 1. Likewise, an area will have a PIV of 5 when it contains only indicator species as- Table 8. Prevalent species list for 10 study sites along with average percent cover in the different areas sampled. Importance _ Study site Species value 1° 3 2 4 5 6 7 8 10 9 Eriogonum corymbosum 1421.5 30.8 13.2 13.6 17.7 13.9 11.5 11.9 9.1 13.3 7.4 Stipa comata 563.1 .2 .4 12.5 1.5 50.9 28.6 Oryzopsis hymenoides 389.5 .3 .7 .4 2.7 6.4 1.2 6.9 18.8 1.2 2.4 Agropyron sniithii 380.8 14.7 38.0 23.3 Artemisia tridentata 285.9 1.3 2 3.5 4.4 3.2 14.3 13.9 Agropyron trachycaulum 202.8 .2 10.4 39.5 .6 Xanthocephalum sarothrae 113.3 .9 13.3 Bromus tectorum 106.0 .4 .1 2.6 9.4 .5 Gilia leptomeria 55.8 2.8 1.3 .7 6.5 Sphaferalcea coccinea 42.7 .9 .4 4.9 Atriplex confertifolia 39.3 2.1 .9 4.3 .6 Chenopodium leptophyllum 37.0 3.2 2.4 .9 2 .2 .5 .2 Chrvsothamnus viscidiflorus 33.2 1.3 .2 .2 .6 .7 14.0 1.2 Cercocarpos montanus 32.7 9.6 6.8 Astragalus tenellus 25.7 8.6 Amelanchier utahensis 23.1 .8 .6 .9 2 .9 Lepidium densiflorum 22.7 4.1 3.5 Agrophron spicatum 21.8 1.2 4.1 8.9 .8 Chrysothamnus nauseosus 21.6 2.3 2.9 Chenopodium fremontii 21.6 3.6 Artemisia nova 21.1 10.9 Lappula redowskii 19.8 1.1 .1 .7 1.5 Sitanion hystrix 18.8 .1 .1 .2 .3 1.7 Poa sandbergii 18.8 1.1 .7 1.1 3.5 Opuntia polyacantha 18.0 .6 .2 3.0 2.9 •Numbers equivalent to study sites as listed in Table 7. 186 Great Basin Naturalist Vol. 39, No. 2 signed adaptation numbers of 5. Areas which contain a mixture of species having different adaptation numbers will have PIV's inter- mediate between 1 and 5. The ordination (Fig. 2) was prepared by placing the computed PIV's so that each area exhibited a linear relationship to all the oth- ers. This ordination (Fig. 2) separates the 10 study sites into two fairly distinct groups. These two groups correlate well with the dis- tributional patterns of the two varieties of Eriogonwn corymbosum discussed earlier. Va- riety corymbosiim occurs in the group to the left, which contains the Mowry Shale study site, the Moenkopi formation study site, the Evacuation Creek and Parachute Creek members of the Green River formation study sites, and the Wells Draw study site. Variety erectum occurs in the group to the right, which contains the Rock Creek study site, the Uinta River study site, the Yellowstone River study site, the Strawberry Valley study site, and the Red Creek study site. By consulting Table 4 and Figure 2, we see that those study sites in the eastern and southern part of the basin are much more alike and become less so as you travel west and northward across the basin. Similarly, those study sites in the north and western part of the basin exhibit high indexes of similarity, but as you travel south and eastward across the basin, the sites become less and less alike. Those sites in the eastern and southern parts of the basin oc- cupy areas which belong to the deserts of the Uinta Basin and are below 5500 feet eleva- tion. Those sites in the north and western part of the basin are all above 6000 feet and are either in or very near the Uinta Moun- tains. This factor would tend to place these sites in areas of higher rainfall, and thus in the nondesert areas of the basin. The response of the prevalent plant species to the ordination is shown in Table 8. Since the patterns of these species along the ordi- nation do not, in themselves, suggest reasons for or factors involved in their distributional patterns, other ecological data collected dur- ing this study were also plotted against the above-described ordination (Figs. 3, 4, 5). This was to determine if any correlation be- tween these factors and distributional pat- terns might be discovered. Total cover and soil depth show positive correlation to the ordination; bare ground, cation exchange capacity, and soluble salts are negatively correlated to the ordination and thus may so influence the distributional patterns of those species which showed some correlation with the ordination. The species are Eriogonwn corymbosum, Chenopodium leptophyllum, Ariplex confertifoUa, Agropy- ron smithii, Artemisia tridentata, Stipa com- ata, Tetradymia spinosa, Agropyron tra- chycaulum, Agropyron spicatum, Cercocarpus montanus, and Xanthocephalum sarothrae. As is evident from studying Table 8, the above species show definite patterns of distribution along the ordination and these patterns ap- pear restricted to certain areas of the ordina- tion. These species, then, can to some extent be classed as indicators of the areas to which they appear restricted. The species E. corymbosum, C. leptophyl- lum, and A. confertifoUa show decreasing composition percents from left to right (Table 8), but the composition of A. smithii, A. tridentata, and S. comata increase from left to right (Table 8). Looking at Figures 3 and 4, we see that shrub cover, bare ground, cation exchange capacity, and total soluble salts decrease from left to right, but total cover, grass cover, diversity, and soil depth increase. This would indicate generally that E. corymbosum, C. leptophyllum, and A. con- fertifoUa are best adapted to areas having low diversity, shallow soil, elevated levels of exchangable ions and soluble salts, and little total ground cover. A. smithii, A. tridentata, and S. comata, on the other hand, grow best in areas of high diversity where the soils are moderately deep and low in exchangeable ions and soluble salts and exhibit high degrees of total ground cover. It is evident that there exist definite rela- tionships between these ecological factors and the distributional patterns of the listed species. From Table 2 and Figures 3, 4, and 5, we see that where total soluble salts and cation exchange capacities are at their peaks and soil depth and total cover are at their lower levels, the plants E. corymbosum, C. leptophyllum, and A. confertifoUa reach their greatest importance in the community. In the case of A. confertifoUa, it occurs only in sites which have appreciable amounts of soluble June 1979 Brotherson, Brotherson: Plant Community 187 Ordinal ion Values Fig. 3. The relationships of community characteristics (i.e., total cover, diversity, shrub cover, and grass cover) to the ordination. Correlation data are shown. = ~198 6x +1038 2 -. 100. a. Y = 47 22 -4.9 + 31.4 f- ;: 60. J- . 20. ~~' ■ ■■ " ■ -r- Ordination Values Fig. 4. The relationships of site characteristics (i.e., soil depth, soluble salts, exposed bare ground, and change capacity) to the ordination. Correlation data are shown. 188 Great Basin Naturalist Vol. 39, No. 2 salts in the soil. Upon examination of the pat- terns shown by A. tridentata, A. smithii, and S. comata, we see that they reach their great- est importance where the reverse of the above conditions are true. Indications are that these last three species are able to with- stand the competition of other plants more readily than the previous three species. On further examination of Table 8, another interesting relationship is made apparent. The three species of Agropyron, A. smithii, A. spicatum, and A. trachycaulum show definite areas of preference or distribution. In each case, the distribution patterns are discrete and show little or no overlap. Of the other factors examined (i.e., percent sand, silt, clay, pH, and calcium, magnesium, potassium, and sodium), none appeared to have any relationship to the distributional patterns of the species studied. But in some cases, a few of the factors did show distinct relationships to other related soil factors. As the percent sand in the soil decreases, the percent of silt increases. The percent clay ap- pears to show some relationship to the above two factors in that it tends to act somewhat like the silt in relationship to the sand. Cal- cium and magnesium also show direct corre- lation (Table 2). As the concentrations of cal- cium decrease or increase, there is a corresponding increase or decrease in the concentrations of magnesium. The concentra- tions of potassium and sodium also appeared to be related (Table 2) in that where the lev- els of sodium are high, the levels of potassium are low and vice versa. Family Composition A total of 173 species of vascular plants, representing a total of 93 genera and 40 fam- ilies, were collected from the above de- scribed study areas. Of these species, 72.7 percent belonged to the families shown in Table 9. The figures indicate that these nine fami- lies contribute about 70 percent of the spe- cies to the cold temperate desert shrub re- l^^:. 100. y=81.- 24 ^80. a • ^„^'^^ > c5-.o. ^,^>^^ £ ..j-'-''''^^ 340. c o 220. • , .>^ . 1 Bare Ground 00. y=794.+240 ^^ 80. • .^^^"'^ 60. • ■^ 40. JT-^^ , 20. • 1 1 100. r=- r2 = 80 64 75. 4.7 2 ^80. o (J 60. ^ . • e i40. Z20. • ." --- • • •"^^ • 1 1 Soil Depth ( dm ) Total Cover Fig. 5. The relations of Eriogontim cover to the ordination and to other site characteristics. Correlation data are shown. 1979 Brotherson, Brotherson: Plant Community 189 gions of the Uinta Basin, regardless of topography, soil depth, soil texture, soil pH, or other factors measured in this study. A similar family composition chart (Table 10) has been computed on an individual study site basis. As can be seen, the nine plant families, with the exceptions of Polygonaceae and Chenopodiaceae, and then these only in two of the study areas, show a fair similarity to the earlier figures. The ecological or phyto- geographical significance of the dominance of these nine families is not known, but fur- ther investigation along such lines tends to hold interest. Discussion There appears in this study evidence that the distributional patterns of some species na- tive to the Uinta Basin are related to measur- Table 9. Dominant plant family composition of 10 study sites. Family Percent Asteraceae Poaceae Brassicaceae Fabaceae Chenopodiaceae Scrophulariaceae Boraginaceae Polygonaceae Polemoniaceae 22.0 9.3 8.7 5.8 5.8 5.7 5.7 5.7 4.0 Total 72.7 able characteristics of their communities. Eriogonum corymbosum, Chenopodium lepto- phyllum, Atriplex con ferti folia, Stipa comata, Artemisia tridentata, and Agropyron smithii are affected by both vegetational and eda- phic factors of the community. Being aware that organisms can and do modify the phys- ical environment (Goiger 1957, Mcintosh 1957, Polunin 1960), it is still apparent that factors which restrict one species in one com- munity may very well allow other species to reach their greatest importance in another environment. The vegetation of the different areas stud- ied showed remarkable similarity at the fam- ily level and extreme variability at the spe- cies level. There was a definite increase in total vegetative cover, grass cover, and diver- sity as you moved from southeast to north- west across the Uinta Basin. As the vegetative cover increased, the soil depth also generally increased. Paralleling these trends, the importance of such plants as Eriogonum corymbosum, Che- nopodiimi leptophylliim, and Atriplex confer- tifolia in the community decreased, while such plants as Stipa comata, Agropyron smithii, and Artemisia tridentata became more important. Other species (i.e., Gilia lep- tomeria, Oryzopsis hymenoides, Chryso- thamnus viscidiflorus, and Amelanchier uta- hensis) did not seem to be affected by such trends. The species (Table 8) Tetradymia spinose, Agropyron trachycauliim, Cerco- carpus montanus, Agropyron spicatum, and Table 10. Comparisons of the total number of plant families, total number of plant species and the percentage of species contributed to the flora by nine major plant families. Total number families Total number _ species Percent of species in nine plant families Study Site 1° 2 3 4 5 6 7 8 9 Mowry Parachute Creek 11 8 29 19 17 37 10 16 5 - 28 5 3 5 10 16 3 16 Evacuation Creek 12 26 27 12 4 — 19 — 4 8 4 Moenkopi Wells Draw 19 18 39 51 18 26 13 12 8 8 5 2 8 12 2 8 4 8 • 6 3 6 Red Creek 15 37 27 8 8 8 8 8 3 8 — Rock Creek 14 35 23 17 3 6 3 6 3 3 3 Strawberry Valley Uinta River 15 16 45 43 24 26 13 12 2 5 11 9 4 5 13 7 2 7 7 5 4 5 Yellowstone River 16 43 23 12 5 - 2 2 7 14 5 °1 = Asteraceae; 2 = Poaceae; 3 = Brassicaceae; 4 = Fabaceae; 5 = Chenopodiaceae; = Polemoniaceae. Scrophulariaceae; 7 = Boraginaceae; 8 = Polygonaceae; 9 190 Great Basin Naturalist Vol. 39, No. 2 Xanthocephalum sarothrae showed definite correlation to certain study sites, but the basis for such correlation was not deter- mined. Of the edaphic factors measured, soil depth, soluble salts, and cation exchange ca- pacity showed positive relationships to the distribution patterns of the above-mentioned species. The distributional patterns of Arte- misia tridentata and Atriplex confertifolia were found to be in agreement with Kearney et al. (1914), Fautin (1946), Thatcher (1959), and Robertson et al. (1966). A. tridentata oc- curred near the mountains on soils moder- ately deep to deep exhibiting fairly low levels of soluble salts. A. confertifolia was found only where the soils showed fairly high levels of soluble salts and where the soils were gen- erally shallow. Paralleling A. tridentata are the distributional patterns of Agropijron smithii and Stipa coniata. The distribution patterns of Chenopodium leptophyllum and Eriogonum corymbosum show some relation- ship to that of A. confertifolia. Factors of the soil, such as pH, concentra- tions of the cations calcium, magnesium, po- tassium, and sodium, and soil texture did not appear to influence the distributional pat- terns of species associated with this study. Eriogonum corymbosum var. corymbosum showed correlation with the above-men- tioned factors as well as with total cover. As is evident from the data, E. corymbosum var. corymbosum prefers areas of low diversity, high shrub cover, shallow soils, high soluble salts, high cation exchange capacities, and low total ground cover. Eriogonum corymbo- sum var. erectum, on the other hand, reaches its highest development in communities of high diversity, high grass cover, and deep soils which are low in soluble salts and cation exchange capacities. That both are found on a number of different geological formations supports the hypothesis that this species is an indicator of peculiar soil types. Other factors involved in the distribution of the above species are competition and community disturbance and erosion. It is sug- gested that competition between species is probably important because field work in- dicates, for example, that as the total cover of a community increases, the importance of E. corymbosum decreases. Although no mea- surements were made on community disturb- ance and erosion, 8 of the 10 study sites showed varying degrees of disturbance and erosion. The 2 sites which did not show ero- sion but only slight disturbance, indicated by the presence of such plants as Bromus tecto- rum and Xanthocephalum sarothrae, also ex- hibited the lowest composition percents for species such as E. corymbosum. It is suggested that E. corymbosum var. co- rymbosum, which is found only in study sites located in the desert areas of the basin, is a shallow soil variety and one that prefers com- munities that show high degrees of disturb- ance, little competition, and fairly high levels of soluble salts in the soil and are found at elevations below 5500 feet. Eriogonym co- rymbosum var. erectum, on the other hand, appears to compete best in communities above 6000 feet that show less disturbance than the desert areas and have deeper soils and low levels of soluble salts. Although the suggested associations be- tween the site characteristics measured in this study and the vegetation do not prove causal relationships, they do serve to aid in the assignment of probable causes of the ob- served vegetational patterns. To gain a com- plete and final insight into the ecology of Eriogonum corymbosum and its associated species of the cold temperate desert shrub re- gions of the Uinta Basin, further and more in- tense investigations of those factors studied in this paper, as well as further investigation of other important community characteristics, will be necessary. Literature Cited Billings, W. D. 1949. The shadscale vegetation zone of Nevada and eastern California in relation to cli- mate and soils. Amer. Midi. Nat. 42:87-102. 1950. Vegetation and plant growth as affected by chemically altered rocks in the western Great Ba- sin. Ecology 31:62-74. 1951. Vegetational zonation in the Great Basin of western North America, pp. 101-122. In Les Bases Ecologiques de la Vegeudration de la Vege- tation de Zones Arides. U. I. S. B. Paris. 1952. The environmental complex in relation to plant growth and distribution. Quart. Rev. Biol. 27:251-265. BouYoucos, G. J. 1936. Directions for making mechani- cal analysis of soils bv the Hydrometer Method. Soil Sci. 42: 225-230. June 1979 Brotherson, Brotherson: Plant Community 191 1951. A recalibration of the Hydrometer Method for making mechanical analysis of soils. Agron. Jour. 43:434-438. Cain, S. A., and G. M. de Oliveira Castro. 1959. Man- ual of vegetational analysis. Harper and Brothers, Pubs., New York. Cannon, H. L. 1952. The effect of uranium-vanadium deposits on the vegetation of the Colorado Plateau. Amer. Jour. Sci. 250:735-770. Dastrup, B. C. 1963. Vegetational changes of the Uinta Basin since settlement. Unpublished master's thesis. Brigham Young University, Provo, Utah. Dix, R. L., AND J. E. Butler. 1960. A phytosociological study of a small prairie in Wisconsin. Ecology 41:316-327. Flowers, S. 1960. Ecological studies of the flora and fauna of Flaming Gorge Reservoir Basin, Utah and Wyoming. Anthropological Papers No. 48, Department of Anthropology, University of Utah. Fautin, R. W. 1946. Biotic communities of the Northern Desert Shmb. ecol. Mono. 16:251-310. Gates, D. H., L. A. Stoddart, and C. W. Cook. 1956. Soil as a factor influencing plant distribution on the Salt-Deserts of Utah. Ecol. Mono., 26:155-175. Geiger, R. 1957. The climate near the ground. Howard University Press, Cambridge, Massachusetts. Graham, E. 1937. Botanical studies in the Uinta Basin of Utah and Colorado. Ann. Carnegie Mus., Pitts- burgh 26:1-432. 1903. Contributions to western botany. Contr. West. Bot. 11:14. Kearney, T. H., et al. 1914. Indicator significance of vegetation in Tooele Valley, Utah. J. Agr. Res. 7:365-417. (Crucheberg, a. R. 1951. Intra-specific variability in the response of certain native plants to serpentine soil. Amer. J. Bot. 38:408-418. 1954. The ecology of serpentine soils III. Plant species in relation to serpentine soils. Ecology 35:267-274. VIarsell, R. E. 1964. Geomorphology of the Uinta Ba- sin—a brief sketch. Guidebook to the geology and mineral resources of the Uinta Basin, Publisher Press, Salt Lake City, Utah. McConkie, D. L. 1941. An economic survey of the Uinta Basin. Unpublished master's thesis. Brig- ham Young University, Provo, Utah. Mc:Intosh, R. P. 1957. The York Woods, a case history of forest succession in southwestern Wisconsin. Ecology 38:29-37. Osmond, J. C. 1964. Tectonic history of the Uinta Basin, Utah. Guidebook to the geology and mineral re- sources of the Uinta Basin, Utah. Publisher Press, Salt Lake City, Utah. Peterson, P. P. 1952. Geology of the Thistle area, Utah. Unpublished thesis. Brigham Young University, Provo, Utah. PoLUNiN, N. 1960. Introduction to plant geography and some related sciences. McGraw-Hill Book Com- pany, Inc., New York. Reveal, J. L. 1967. Notes on Eriogonum V: A revision of the Eriogonum corymbosum complex. Great Ba- sin Nat. 27:183-229. Robertson, D. R., et al. 1966. Vegetation and soils of al- kali sagebrush and adjacent big sagebrush ranges in North Park, Colorado. J. Range Manage. 19:17-20. Russell, D. A. 1948. A laboratory manual for soil ferti- lity students, 3d ed. Wm. Brown Company, Du- buque, Iowa. Shantz, H. L., and R. L. Piemeisel. 1940. Types of veg- etation in Escalante Valley, Utah, as indicators of soil conditions. U.S. Dept. Agric. Tech. Bull. 713. SoRENSEN, T. 1948. A method of establishing groups of equal amplitude in plant sociology based on sim- ilarity of species content. Konge, Dan. Vidensk. Selsk. JS:4:l-34. Thatcher, A. P. 1959. Distribution of sagebrush as re- lated to site differences in Albany County, Wyoming. J. Range Manage. 12:55-61. Walker, R. B. 1954. The ecology of serpentine soils II. Factors affecting plant growth on serpentine soils. Ecology 35:259-266. Welsh, S. L. 1957. An ecological survey of the vegeta- tion of the Dinosaur National Monument, Utah. Unpublished thesis. Brigham Young University, Provo, Utah. Whittaker, R. H. 1954. The ecology of serpentine soils IV. The vegetation response to serpentine soils. Ecology 35:275-288. VARIATION IN HEMOGLOBIN TYPES IN THE DEER MOUSE (PEROMYSCUS MANICULATUS) ALONG AN ALTITUDINAL GRADIENT David Wasserman' and Donald J. Nash' Abstract.— Deer mice (Peramyscus manictikittis) were captured along an altitudinal gradient that extended from 5,000 feet (1524 m) up to 11,000 feet (3353 m) in central Colorado during August and early September, 1976. Starch gel electrophoresis of deer mouse hemoglobin followed no clear trend that would indicate that slight biochemical differences in the molecule help facilitate adaptation to the decreased p02 that exists at that altitude. Organisms that Hve at high altitudes must in some way adapt to the hypoxic conditions and meet their oxygen needs. Gluecksohn- Waelsch (1960) suggested that multiple he- moglobins may differ in physical properties and thus facilitate environmental adapta- tions. In one known mutant human hemoglo- bin, hemoglobin Rainier, on which a histidine residue replaces a tryosine residue at one point, oxygen affinity is greatly enhanced (Stamatoyannopoulos et al., 1968). Studies by Ahl (1968) and Sawin (1970) looked at the electrophoretic patterns found in deer mouse hemoglobin along an altitudinal gradient. In both of these studies a predominance of dif- fuse (double)-banded individuals were found at low altitude. High altitude groups were characterized by a predominance of single- banded individuals. If the hypothesis put forth by Gluecksohn-Waelsch (1960) is cor- rect, then the patterns discovered in the ear- lier studies should be found along other al- titudinal clines if these biochemical differences in the hemoglobin molecule facil- itate greater oxygen-binding efficiency. Methods A total of 67 Peromyscus maniculatus were trapped from 27 July through 8 September 1976 in Larimer and Pitkin counties, Colo- rado, at altitudes of 5,000 feet, 6,500 feet, 8,000 feet, and 11,000 feet. Two populations were sampled at each altitude. Blood samples were collected from the or- bital sinus using heparinized capillary tubes. The blood was centrifuged immediately, and both cells and plasma were frozen with dry ice and later transferred to a freezer and stored at -20 C. Hemoglobin was analyzed by vertical starch gel electrophoresis, using the method of Smith (1968). Following electrophoresis, gels were sliced, fixed, and stained with bro- mophenol blue. Results and Discussion A number of studies have dealt with the electrophoretic variants in both the serum and cellular fractions of the blood of the deer mouse. Most studies demonstrated that Per- omyscus maniculatus has at least two elec- trophoretically separable hemoglobins (Fore- man 1960, Ahl 1968, Sawin 1970). Studies of the genus Peromyscus showed that there are variations in the double-banded phenotypes (Rasmussen 1970, Selander et al. 1971). A third hemoglobin variant was reported in one wild mouse captured near Flagstaff, Arizona, in 1967 (McCracken and Foreman 1971). Re- cent papers have verified the existence of the third variant and three triple-banded pheno- types (Jensen et al. 1976, Maybank and Daw- son 1976). It should be noted that the triple- banded individuals were either Foreman's original stock or low altitude populations (600-4700 ft.). 'Department of Zoology and Entomology, Colorado State University, Fort Collins, Colorado 80523. 192 fune 1979 Wassehman, Nash: Deeh Mouse Hemoglobin 193 In previous studies, low altitude deer mice were shown to either have a predominance of individuals exliibiting the diffuse pheno- type (type R) or only the diffuse phenotype (Ahl 1968, Sawin 1970). At high altitudes (above 7,000 feet) the single-banded pheno- type (type S) was the predominant variety found (Ahl 1968, Sawin 1970). Results of the present investigations do not agree with the previous studies; however, the sampling tech- niques made it possible to ascertain if the variation found was between altitudes and/ or intraaltitudinal in character. In the populations that were sampled, there was no discernible change in frequen- cies of the phenotypes over the 6,000 ft. range (Table 1). Population A (5,000 ft. sample) had three individuals that exhibited the single-banded phenotype, but all other individuals in the 5,000 ft. group exhibited the diffuse phenotype. Throughout the re- mainder of the populations sampled, all indi- viduals exhibited the diffuse phenotype ex- cept for two individuals in population I (11,000 feet) that exhibited a triple-banded variety of hemoglobin. It was not possible to ascertain which of the three triple-banded phenotypes as outlined by Jensen et al. (1976) and Maybank and Dawson (1976) was pres- ent. It appears to be the first report of the ex- istence of triple-banded hemoglobin pheno- types in high altitude populations. In light of these findings, the hypothesis put forth by Gleucksohn-Waelsch (1960), which suggests that multiple hemoglobins may differ in physical properties and thus fa- cilitate environmental adaptations, must be questioned. While the two studies noted above found a high frequency of single-band- ed individuals at higher altitudes, the present results do not follow a similar pattern. The genetic polymorphisms found along altitudinal gradients could be a product of genetic drift or varying selective pressures in different environments. The findings suggest that the structural variations found in the beta chain of the hemoglobin molecule in P. manicidatits do not infer a greater ability on the part of the organism to exist at high alti- tudes. At the present time it is not known if these slight structural differences manifest themselves in any manner in the deer mouse. The hypothesis put forth by Gleucksohn- Waelsch (1960) could be tested in an alter- nate fashion by comparing the 02-binding ca- pacities of equal volumes of blood of each phenotype. Literature Cited Ahl, a. S. 1968. Electrophoretic examination of he- moglobin and plasma proteins from three alti- tude groups of Pewmyscus manicttlatus nebras- censis. Comp. Biochem. Physiol. 24:427-435. Foreman, C. W. 1960. Electromigration properties of mammalian hemoglobins as taxonomic criteria. Amer. Mid. Nat. 64:177-186. Gluecksohn-Waelsch, S. 1960. The inheritance of he- moglobin types and other biochemical traits in mammals. J. Cell. Comp. Phys., 56:89-101. Jensen, J. N., J. R. Merkle, and D. I. Rasmussen. 1976. Locus number and multiple hemoglobins in Eu- tamias and Peromyscus. Biochemical Genetics 14:541-545. Maybank, K. M., and W. D. Dawson. 1976. Genetic and developmental variation of hemoglobin in the deer mouse, Peromyscus manictdatus. Bio- chemical Genetics 14:389-400. Table 1. Hemoglobin variants and frequencies at trapping sites within altitudes. Altitude in feet Population Type S Type R Type F Total 5000 6500 8000 9500 11000 3 (.50) 3 (.50) 7(1.0) 5(1.0) 6 (1.0) 6 (1.0) 9 (1.0) 8 (1.0) 7(1.0) 6(1.0) 5 (.70) 2 (.30) 194 Great Basin Naturalist Vol. 39, No. 2 McCracken, K. M., and C. W. Foreman. 1971. A new polymorphism and systematics in Peromyscus I hemoglobin variant in the deer mouse (Per- Variation in the old field mouse {Peromyscus po- amysciis maniculatus). ASB Bui. 18:45. lionotus). Stud. Genet. VI. Univ. Texas Publ. Rasmussen, D. I. 1970. Biochemical polymorphisms and 7103:49. genetic structure in populations of Peromyscus. Smith, I. (ed.). 1968. Chromatographic and Electro- Symp. Soc. Lond. 26:.335-348. phoretic Techniques. Vol. II. Zone Electro- Sawin, C. F. 1970. The respiratory function of the blood phoresis. Inter-science Publishers, New York. 524 of deer mice at sea level and high. Unpublished p- dissertation. University Microfilms, Ann Arbor. Stamatoyannopoulos, G., A. Yoshida, J. Adamson, Selander, R. K., M. H. Smith, S. Y. Yang, W. E. and S. Heinenberg. 1968. Hemoglobin with in- Johnson, and J. B. Gentry. 1971. Biochemical creased O2 affinity. Science 158:712-713. FIRST RECORD OF PATAPIUS SPINOSUS IN IDAHO AND NEVADA (HEMIPTERA: LEPTOPODIDAE) Donald R. Brothers' Abstract.— The first record of Patapitis spinosus (Rossi) in Idaho and Nevada is reported, having previously been found in America onlv in California. The family Leptopodidae is represented in America by a single introduced Eastern Hemisphere species, Patapius spinosus (Ros- si). This species was first reported in America in the United States by Usinger (1941) from a single specimen collected from Colusa Coim- ty, California. Since that time, it has been collected in a number of California counties in increasing numbers (Drake 1954). To my knowledge, there are no published United States records of P. spinosus occurring out- side of California. Recent collections of P. spinosus from Idaho and Nevada establish the first state record for this insect and repre- sented an eastern extension of its known range. Collection records are as follows: Idaho: Gem Co., Pearl vicinity, T6N,R1E,S14, 4400', IX-9-77, IX-14-77, IX-26-77, X-2-77 (D. R. Brothers) (G. A. Shook), adults and nymphs; near Payette River, 12 miles E Emmett, T7N,R1E,S24, 2580', iV-14-78 (G. A. Shook), adults. Nevada: Reno, 1971 (M. A. Bechtel). Although Woodward et al. (1970) reported that leptopodids frequent the drier parts of rocks in streams, it appears that this species may not be restricted to that habitat. Usin- ger's specimen was found under a piece of fibrous tree protection material on the trunk of an almond tree. Idaho specimens from the Payette River area were taken on the under- side of a board in a cottonwood (Populus) grove adjacent to the river. Adults and nymphs taken from the vicinity of Pearl were found on the underside of cobble-sized rocks in a mine tailings dump in an area dominated by sagebrush {Artemisia) and rabbitbrush {Chrysothanums). The nearest known per- manent water from this site is approximately one mile to the west. Adjacent to the site. however, is an intermittent stream which has water for a few months during spring runoff. Since P. spinosus has now been collected east of the Sierras in habitats common to many parts of the Pacific Northwest and Great Basin, it can be anticipated that addi- tional collections of this insect will be made in these areas. Collectors should watch for a small (3-4 mm in length) saldidlike bug with long spines on much of its body, including the eyes (Fig. 1). Idaho specimens of P. spinosus are depos- ited in the California Academy of Science, San Francisco, California; U.S. Natural Mu- seum, Washington, D.C.; Brigham Young University, Provo, Utah; University of Idaho, Moscow, Idaho; and Boise State University, Boise, Idaho. Acknowledgments Appreciation is expressed to Dr. Jon L. Herring of the U.S. National Museum for confirming my identification of Idaho speci- mens and allowing me to report the Nevada record, and to Dr. Charles W. Baker of Boise State University for critically reading the manuscript. Literature Cited Drake, C. J. 1954. An undescribed leptopodid from In- dia (Hemiptera). J. Kansas Ent. Soc. 27:111-112. Usinger, R. L. 1941. A remarkable immigrant leptopo- did in California. Bull. Brooklyn Ent. Soc. 36:164-165. 'Woodward, T. E., J. W. Evans, and V. F. Eastop. 1970. Hemiptera. p. 441. In: Commonwealth Sci- entific and Industrial Research Organization. 1970. The insects of Australia. Melbourne Uni- versity Press, Melbourne, xiii + 1029 pp. 'P.O. Box 8413, Boise, Idaho 83707. 195 196 Great Basin Naturalist Vol. 39, No. 2 mm Fig. 1. Patapius spino.ms, dorsal view. BEE VISITATION OF PHLOX BRYOIDES (POLEMONIACEAE) V. J. Tepedino^'^ Abstract.— Collections of diurnal insects from a population of Phlox hnjoides on shortgrass prairie in southeastern Wyoming showed the flowers to be visited predominantly by Synhalonia fulvitarsis, a long-tongued bee. Analysis of pollen loads carried by captured bees revealed that almost all individuals were collecting Phlox pollen. P. bryoides may be an exception to the generalization that the genus Phlox is exclusively pollinated by Lepidoptera. Entomophilous pollination in the genus Phlox is thought to be effected almost exclusi- vely by Lepidoptera (Grant and Grant 1965). Species with corollas 1-2 cm long are ex- clusively pollinated by perching butterflies; those with corollas 3-4 cm long are polli- nated by hawkmoths (Grant and Grant 1965). Based upon observations of three species (P. caespitosa Nutt., P. diffusa Benth., P. multi- flora A. Nels.) plus a citation for P. andicola (Britt.) (Wherry 1955), it was concluded that all species of the Section Occidentales ( = Section Microphlox of Wherry 1955), which "contains over 20 species of woody- based, needle-leaved, cespitose or cushion- like shrubs," were pollinated by Lepidoptera (Grant and Grant 1965). In particular, west- em cushion plants with erect 1 -cm-long co- rolla tubes are pollinated by perching noc- tuid moths (Grant and Grant 1965). However, at least two species in the Section Occidentales possess corollas that are sub- stantially shorter than 1 cm (P. bryoides Nutt., P. hoodii Rich., measurements from Wherry 1955) and could be pollinated by long-tongued bees or flies. Indeed, data re- ported here from a population of P. bryoides, a dense cushion plant common on shallow soils and rocky outcrops in southeastern Wyoming, suggest that the Grants' general conclusion (1965) may require modification. Individuals of the long-tongued eucerine bee species Synhalonia fulvitarsis fulvitarsis (Cresson) were common pollen collectors on the flowers and may be important pollina- tors. Phlox bryoides blooms from the last half of May to late June/early July, with peak flow- ering during the first half of June. During this period in 1975 and 1976, weekly collections of flower-visiting bees were made during four hour-long periods, two each in morning and afternoon, on an 11000-m- plot in Albany Co., 8 km SSE of Laramie. About 30 percent of the plot was covered heavily by P. bryoides. Collections were not restricted to P. bryoides; rather, each floral species was cen- sused in rough proportion to its percent of total floral abundance during each period. Thus the number of insects visiting P. bryoides was much larger than the relatively small numbers reported here. Synhalonia fulvitarsis works the salver- form flowers of P. bryoides very rapidly, vis- iting only a few of the many available in each cvishion before moving to another cush- ion. In 1975, 11 females, 10 carrying pollen, were captured on the flowers of P. bryoides; in 1976 7 of 8 females captured had Phlox pollen loads. No diurnal pattern in foraging was evident except that females did not begin visiting flowers until after 1030 hours. There- after, activity was evenly spaced until collec- tion ended, generally between 1530 and 1600 hours. Other bee species found on Phlox were represented by only 1 or 2 individuals each, and they did not carry pollen. Diurnal Lepi- doptera were rare on the flowers. 'Department of Zoology and Physiology, University of Wyoming, Laramie, Wyoming S 'Present address: Bee Biology and Systematics Laboratory, Agricultural Research, Scii UMC 53, Logan, Utah 84322. ■ and Education Administration, USDA, Utah State University, 197 198 Great Basin Naturalist Vol. 39, No. 2 Most pollen loads examined from the 17 specimens (both years combined) contained more than 70 percent Phlox pollen (Fig. 1). Indeed, seven females carried loads that were almost pure (90 percent Phlox or higher), and the mean for all pollen loads was 64.5 per- cent Phlox (S. D. = 35.4, range 5.6-100.0). Although Timberlake (1969) also reported S. fiilvitarsis from the flowers of Phlox, this association does not appear to be obligatory. Rather S. fiilvitarsis is probably polylectic, because it has been recorded from a large number of plant species (Timberlake 1969). Effective cross-pollination in the Section Occidentales seems to require a nectar rather than pollen-collecting flower visitor because the anthers are situated above the stigma while the nectaries are located at the base of the corolla tube (Grant and Grant 1965). Thus, species that collect only pollen would not contact the stigma. In most species of Phlox the length of the corolla tube precludes nectar collection by all but long-tongued Lepidoptera. In P. bryoides, however, (and in P. hoodii as well) the corolla tube is relatively short (4-8 mm, Wherry 1955) and the nec- taries are well within the reach of long- 30 Percent 70 100 Phlox Pollen Fig. 1. Distribution of pollen loads carried by Sijnha- lonia fiilvitarsis by percent Phlox bryoides. tongued bee species such as S. fiilvitarsis (proboscis length measured from base of mentum through flabellum; x = 8.44, s.d. = 0.465, N = 20? ? ). It is not unlikely that S. fiilvitarsis collects nectar from P. bryoides at the same time pollen is collected, thereby achieving pollination. Whether P. bryoides is dependent upon long-tongued bees for pollination in any consistent way awaits the examinations of other populations. Although bee pollination is prevalent in the family Polemoniaceae, it is regarded as the primitive condition from which all other pollination systems have been derived (Grant and Grant 1965). Moreover, the genus Phlox is regarded as an advanced member of the family, in part because of its pollination syn- dromes (Grant and Grant 1965). Wherry (1955) viewed Microphlox as an advanced section of the genus Phlox on morphological grounds and regarded the subsection Canes- centes, of which P. bryoides and hoodii are members, as perhaps the most advanced group in the Microphlox. If both inter- pretations are correct, then bee pollination in the Canescentes represents reversion to the primitive condition in an advanced branch of the family. Acknowledgments I appreciate the helpful suggestions on the manuscript provided by Verne Grant, Arthui Holmgren, and P. F. Torchio. Literature Cited Grant, V., and K. A. Grant. 1965. Flower poUinatioi in the Phlox family. Columbia University Press New York. Timberlake, P. H. 1969. A contribution to the system atics of North American species of Synlialonk (Hymenoptera, Apoidea). Univ. Calif. Publ. Ento mol. 57:1-76. Wherry, E. T. 1955. The genus Phlox. Morris Arbore tum Monogr. Ill, Philadelphia. ECOLOGICAL DISTRIBUTION OF RODENTS IN CANYONLANDS NATIONAL PARK, UTAH David M. Armstrong' Abstract.— Studies of microhabitat of 14 species of rodents by cluster analysis suggested that the diverse land- scapes of Canyonlands National Park, Utah, include six broad "habitat-types": (1) rimrock; (2) desert shrublands; (3) saxicoline woodland and sagebmsh; (4) oakbrush; (5) riparian deciduous woodland; and (6) grasslands. Perognathus parvus and Neotomu cinerea were the species most strongly associated with single "habitat-types," desert shrub and saxicoline woodland, respectively. Perornyscus maniculatus and P. truei were the species associated with the broadest ranges of habitats. The rodents with the most similar habitats were Neotoma mexicana and Perornyscus boylii; Eu- tmnias quadrivittatus, P. truei, and P. crinitus; Ammospermophilus leucurus and P. maniculatus; and Dipodomys ordii and Onychornys leucogaster. An understanding of ecological distribution of organisms is important for both inter- pretation and management of ecosystems. Typically the vertebrate ecologist describes patterns of ecological distribution in terms of vegetational associations recognized a priori. The purpose of this paper is to allow the ro- dent fauna itself to define salient patterns of environmental features in Canyonlands Na- tional Park and meaningful associations of mammalian species. This not only provides a description of environmental patterns, but al- lows field naturalists to check their sense of the landscape against mammalian habitats, rather than the opposite (i.e., forcing species' distributions into their view of environmental pattern). Canyonlands National Park preserves some 450 square miles (1170 km-) of spectacular canyons and mesas in San Juan and Wayne counties, southeastern Utah. The park in- cludes the confluence of the Green and Colo- rado rivers, which are entrenched in canyons up to 2000 ft. (610 m) deep. These canyons divide the park (and the rest of southeastern Utah) into three distinct land masses. Eleva- tions in the park range from about 3750 to nearly 7000 feet (1150-2135 m). This range of relief dictates a wide variety of physical conditions and a complex distribution of biot- ic communities. Knowledge of mammals of the Canyon- lands is rather scanty. The area was ignored by the exploratory parties that provided fun- damental knowledge of mammalian distribu- tion elsewhere in the West, such as the Rail- road Surveys of the 1850s and the Bureau of Biological Survey in the early 1900s. John Wesley Powell's expeditions of the 1870s paid almost no attention to the biota of the region. Although the U.S.-IBP Desert Biome Project worked over much of the desert Southwest, no study area was located on the Colorado Plateau (MacMahon 1976). The Canyonlands Section of the Colorado Plateau physiographic province is a showcase for the effects of erosion on an arid land dominated by flat-lying sedimentary strata. For details of geology, see Baars (1971) and Lohman (1974). The climate of Canyonlands is arid, with hot summers, cold winters, and pronounced diel fluctuations in temperature. Mean annual precipitation is about 7.5 in., about one-third of which falls during the third quarter of the year, usually as local, tor- rential thunderstorms (Tanner 1965). Excep- ting the immediate vicinity of the master streams, perennial surface water is limited to a few widely scattered springs and seeps. Bare rock comprises more than half of the surface. Where soils have formed, they are reddish, gravelly to silty loams, moderately 'Department of Integrated Studies and University Museum, University of Colorado, Boulder, 80309. 199 200 Great Basin Naturalist Vol. 39, No. 2 alkaline in reaction (Wilson et al. 1975). Un- developed aeolian sands and silts are present locally. Vegetation of the Canyonlands varies widely with physiographic setting, edaphic conditions, available moisture, and grazing history. Hayward et al. (1958) described four principal vegetation types in the vicinity of Arches National Park: (1) cottonwood-wil- low-tamarisk floodplain, (2) northern desert shrub, (3) pinyon-juniper woodland, and (4) hanging gardens. Although dominance changes locally, these types point out associ- ations that are recognizable in the landscape (and are reflected in mammalian distributions to some extent). On relatively stable inter- fluves, thin, silty soils form. Such flats and rockbound parks are clothed with grassland. Oryzopsis, Hilaria, Stipa, Sporobolus, and Botiteloua are important genera of grasses; Yucca, Opuntia, Gutierrezia, and a variety of annual forbs are present also. Areas of rim- rock, slickrock, and canyon walls are a fre- quent topographic type. These areas often are precipitous; typical substrate is a coarse, unstable coUuvial rubble. Vegetation on such sites includes a variety of shrubs, among them Cowania and Shepherdia on slopes, and Ma- honia and Qiiercus at bases of cliffs. Wood- land of juniper or juniper and pinyon occurs locally on such sites and also on well-drained mesa tops. The understory in this community varies, apparently with edaphic conditions. Phreatophytic cottonwoods (Populus) and willows (Salix) or exotic saltcedar (Tamarix) occur along the major washes. Flqodplains support stands of halophytic shrubs {Sarco- batus, Atriplex). Sagebrush (Artemisia) often occurs in association with junipers or as an overstory on grassy flats. "Hanging gardens" develop locally as mesic associations watered by seepage at contacts between some rock units. These associations are comprised of a striking variety of plants, including Mimulus, Aquilegia, Habenaria, and Rhus. Despite generally forbidding physical con- ditions, Canyonlands National Park supports a diverse vertebrate fauna, including some 60 species of mammals. For general information on the region as a whole, see Hayward et al. (1958). Tanner (1965) provided notes on a few species of rodents. Durrant and Dean (1959) commented briefly on ecological dis- tribution of rodents in Glen Canyon (now in- nundated by Lake Powell), immediately south of Canyonlands. Johnson (1976) and Clevenger (1977) have presented data on some aspects of ecology of rodents in Can- yonlands National Park. For further informa- tion on mammals of southeastern Utah, see Benson (1935), Durrant (1952), Durrant and Dean (1959), Kelson (1951), Lee (1960), and Armstrong (1977b, in press). This report concerns 14 species of rodents, most of them abundant and widespread (ver- nacular name, sample size in parentheses): Eutamias quadrivittatus (Colorado chip- munk, 64), Ammospermophilus leucurus (white-tailed antelope squirrel, 23), Per- ognathus apache (Apache pocket mouse, 29), P. parvus (Great Basin pocket mouse, 35), Dipodomys ordii (Ord's kangaroo rat, 88), Reithrodontomys megalotis (western harvest mouse, 24), Peromyscus crinitus (canyon mouse, 124), P. maniculatus (deer mouse, 128), P. boyUi (brush mouse, 82), P. truei (pin- yon mouse, 202), Onychomys leucogaster (northern grasshopper mouse, 49), Neotoma mexicana (Mexican woodrat, 45), N. lepida (desert woodrat, 34), and N. cinerea (bushy- tailed woodrat, 20). Rodents represented by too few specimens for analysis are Spenno- phihis variegatus, Thomomys bottae. Castor canadensis, Neotoma albigula, and Erethizon dorsatum. Methods Field work on mammals of Canyonlands National Park began in 1972 and continued intermittently to 1978, the principal aim being to provide a range of data on natural history basic to a popular account of the fauna for the National Park Service. Given the broad aims of the research program of which this report is a part, data were gath- ered by various means. Whatever the source of a specimen, its habitat was described as the most prominent feature of plant cover within 1 m of the trap. When no plant was within this radius, a physical descriptor of the trapsite was noted. Analysis of data follows the method utilized by Armstrong (1977a). The similarity index used is Fq/F^ + F^, where Pc is the sum of percentage occur- rences in common, and P^ and Pg are per- I June 1979 Armstronc;: Rodent Ec;olo(;ical Distribution 201 centage occurrences of the two descriptors under comparison. Use of relative (rather than absohite) frequency obviates some prob- lems of differences in sample size. Cluster analysis was by the unweighted pair-group method of Sokal and Sneath (1963:309). Specimens collected in the course of this work are housed in the University of Colo- rado Museum. Results and Discussion Analysis of data was designed to answer three kinds of questions: (1) What associ- ations of habitat descriptors have reality to the rodent fauna? (2) How broadly dis- tributed are species across those "habitat types"? (3) What associations of rodents are found in given habitats? An answer to the first question should approximate a "mouse's- eye view" of the mosaic of habitats. The sec- ond question approaches the phenomenon of fidelity; how faithful are rodents to their hab- itat? Reasonable answers here could be quite helpful in making predictive statements about habitat management. Answers to the third question suggest groups of species that may be worthy of further study from the standpoint of niche structure or competitive interactions. Figure 1 is a cluster diagram of 66 descrip- tors of habitat, based on indices of similarity of rodents associated with each descriptor. Overall mean similarity in the matrix upon which this diagram was based was 0.1783. Taking a mean similarity of 0.450 as an arbi- trary cut-off point, there are nine major sub- clusters of descriptors in the diagram. Group I includes descriptors of slickrock and rim- rock areas, including woodrat dens, most of which are beneath rocky rims. Group II in- cludes many descriptors of open shrublands with poorly developed soils and silty blow- outs or dune sand. Group III is quite com- plex; it includes descriptors of juniper wood- land and broken rocky habitats as well as sagebrush stands. Saltbush and tamarisk also appear in this subcluster. Group IV centers around Gambel's oak and represents the rela- tively mesic brushlands common at bases of cliffs in the Cave Springs area of the Needles District. Group V describes phreatophytic cottonwood-willow woodland of major wash- es and canyon bottoms. Group VI includes grasses and forbs typical of open flats. Groups VII, VIII, and IX are closely related neither to each other nor to other subclusters. All represent descriptors with small samples of rodents associated. The closest resemblance of subcluster VII is with group VI; both groups describe grasslands. Groups VIII and IX truly are miscellaneous, although group VIII does include several de- scriptors of relatively mesic cliffside habitats: Cowania, Cercocarpus, Amelanchier, hanging gardens. These subclusters form a complex pattern, not as nearly comformable as one might hope with the sorts of habitat-types that have been described by previous workers (e.g., Hayward et al. 1958), or the units that the field natu- ralist extrapolates from the landscape. One reason for this is the great ecological ampli- tude of the most abundant species in the sample, Peromyscus truei, which is about equally abundant in sagebrush and in juniper stands. Figure 2 indicates the cumulative percent- age distribution of each rodent species with respect to the nine major subclusters identi- fied in Figure 1. This allows a look at the de- gree of fidelity of species to certain environ- mental attributes. First, note that all species have an association with a single subcluster of descriptors of greater than 40 percent; in- deed, all species except R. megalotis, P. ma- niculatus, and P. crinittis show primary asso- ciations of greater than 50 percent. Five species, A. leucurus, P. parvus, D. or- dii, R. megalotis, and O. leucogaster, show a primary association with subcluster II, repre- sentative of open shrublands on silty to sandy soils. Of these, A. leucurus also shows rela- tively strong secondary associations with groups I and III. Most often, antelope ground squirrels occur in the narrow ecotone be- tween rocky situations and desert flats. D. or- dii shows strong secondary association with group III. This is due to its frequent occur- rence in stands of sagebrush. Reithrodon- tornijs megalotis also has a strong secondary association with group III; harvest mice usu- ally are found on floodplains which may have cover of greasewood (Group II), saltbush or tamarisk (group III) or phreatophytic wood- 202 Great Basin Naturalist Vol. 39, No. 2 land (Group V). Onychoniys leticogaster is similar in local distrijjution to D. ordii (also see Fig. 3, beyond). The species most strongly associated with group II is P. parvus, a spe- cies found only in the Maze District. This species is known from a wide variety of habi- tats in Utah (Hayward and Killpack 1958), but does not seem to be particularly eu- ryecious in the park, occurring mostly under sparse cover of blackbnish (but on a variety RESEMBLANCE .600 Fig. 1. Cluster diagram of 66 habitat descriptors, based on similarity of associated species of rodents. Abbrevia- tions: A., Atriplex, Q., Qtierciis (for explanation of index, see text). 1979 Armstronc;: Rouknt Ecx)lo(;k:al Distribution 203 of substrates, from dune sand to cobbly desert pavement). Only P. apache shows a primary association with group VI, which describes grassland. The Apache pocket mouse is con- siderably more stenoecious than its larger congener, P. parvus, being closely restricted to bunchgrass flats on sandy to silty soils. All four species of Peromyscus show their primary association with group III, rocky habitats. Three of the four show strongest secondary association with subcluster II, shrublands; P. boijlii is the exception, with a strong secondary association with oak brush (Group IV). This analysis is sufficiently crude that it tends to make these species look more similar in ecological distribution than they may actually be. It is not at all uncommon to take three or even four species of Peromyscus in adjacent traps, particularly in broken country. Frequently the animals occur in "text-book" fashion: P. crinitus on shckrock, P. nianiculatus in open shrubs, P. boylii be- neath oakbrush, and P. truei with junipers. These relationships are partially obscured in the present analysis by data from locahties at which fewer species co-occur or in which en- vironments are too complex for the methods used. It is a seeming paradox that no species of Neotoma is related strongly with Group I, which includes the descriptor "woodrat dens." This reflects the fact that woodrats are more difficult to trap in the immediate vicin- ity of their dens than on their foraging range away from the den. Figure 2 suggests that N. lepida is the most euryecious of local species of Neotoma, although all species have strong primary associations with subcluster III. Figure 3 is a cluster diagram of similarity indices of 14 species of rodents with respect to descriptors of habitat. Mean resemblance ,100 400 .600 A^ .800 I 1.000 \r \ 1 MEAN RESEMBLANCE /V. cin0raa N.mexicana P. boylii E. quadrivittatus P. truei P. crinitus N lepida A. leucurus P. manicdiatus D. ordii O. leucogaster R. me gal Otis ■ Pparvjs ■ P. apache B Fig. 2. Cumulative percentage distribution of 14 species of rodents with respect to nine subclusters of habitat descriptors. 204 Great Basin Naturalist Vol. 39, No. 2 in the similarity matrix on which the diagram was based is 0.5297. The diagram shows two different "habitat groups" of rodents, one (group A) occupying broken, rocky habitats, the other (group B) restricted to flats with relatively well-developed soils. The strongest associations are between N. mexicana and P. boylii, E. quadrivittatus, P. truei, and P. cri- nitus, A. leucurus and P. maniculatus, and D. ordii and O. leucogaster. Neotoma mexicana and P. boylii co-occur regularly in saxicoline oakbrush and Mahonia thickets in the Nee- dles District. Eutamias quadrivittatus, P. truei, and P. crinitus occur in rough, broken terrain, P. truei most often in scattered juni- per woodland, P. crinitus more frequently in more open situations. Peromyscus manicu- latus and A. leucurus, which are related closely to the saxicolous group, are species that occupy the ecotone between the two broad habitats; they seem to be about equally likely to be captured among rocks or in open country. Dipodomys ordii and O. leucogaster occur in open shrub- or grassland on sandy soils. The pattern of dispersion across groups of descriptors in Figure 2 suggests that P. truei and P. maniculatus are the most eu- ryecious of local rodents. They also are the species with the highest mean habitat sim- ilarity to all other species, 0.702 and 0.672, respectively. Perhaps the most striking feature of the foregoing analyses is the strong microhabitat similarities among the saxicoline rodents. CUMUi-ATIVE PERCENIAGE 40 60 Fig. 3. Cluster analysis of indices of similarity of 14 species of rodents with respect to descriptors of habitat (for explanation of index, see text). June 1979 Armsth().\(;: Rodent Ec:ological Distribution 205 This is especially noteworthy among closely related species of cricetines, Peromyscus and Neotoma. Other criteria by which these spe- cies assort resources to allow coexistence are under study. The only previous study of ecological dis- tribution of rodents in the general vicinity of Canyonlands National Park was that by Hay- ward et al. (1958), who reported on Arches National Monument as one of several study areas. The suite of species considered was slightly different {Thomomys bottae was in- cluded, but neither P. parvus nor N. mexi- cana was) and the approach was geographi- cally broader and more anecdotal. Still, correspondence with results of the present study is close. Hay ward et al. (1958:32, Fig. 16) showed E. qiiadrivittatus, A. leucurus, and P. truei as considerably more stenoecious than they are in Canyonlands. They pointed out that N. lepida is more broadly distributed ecologically than is N. cinerea, a fact sugges- ted by our data. Acknowledgments Several people and organizations have contributed to the study of the mammals of Canyonlands National Park. David May, chief naturalist, encouraged the work and has provided logistic support. Among National Park Service personnel, David W. Johnson, Peggy Johnson, David Harwood, and Walter Loope deserve special thanks for their help in the field. Charles L. Curlee, James G. Owen, S. Scott Panter, and William C. Sears provided field assistance, and Michael John- son and James C. Halfpenny did considerable independent field work as graduate research assistants. My family, Ann, Jack, and Laura, deserve recognition not only for their help with field work but also for consenting to live in the field for several months. Financial sup- port has been provided by the Society of the Sigma Xi (1972), a Summer Research In- itiation Faculty Fellowship from the Council on Research and Creative Work of the Uni- versity of Colorado (1973), the Penrose Fund (Grant 7615) of the American Philosophical Society (1976), and the Colorado State Uni- versity National Park Service Cooperative Studies Unit (1977 and 1978). Literature Cited Akmstronc, D. M. 1977a. Ecological distribution of small mammals in the Upper Williams Fork Ba- sin, Grand County, Colorado. Southw. Nat. 22:289-304. 1977li. Distributional patterns of mammals in Utah. Great Basin Nat. .37:457-474. in press. A distributional checklist of rodents of Canyonlands National Park, Utah. Occas. Papers Mus., Texas Tech Univ. Baars, D. L. 1972. Red rock country: the geologic his- tory of the Colorado Plateau. Doubleday/Natural History Press, Garden City, 264 pp. Benson, S. B. 1935. A biological reconnaissance of Nav- ajo Mountain, Utah. Univ. California Publ. Zool. 40:439-455. Clevenger, G. a. 1977. The effect of campgrounds on small mammals of Canyonlands and Arches Na- tional Parks, Utah. Unpublished thesis, Utah State Univ., Logan, viii -I- 53 pp. DuRRANT, S. D. 1952. Mammals of Utah, taxonomy and distribution. Univ. Kansas Publ., Mus. Nat. Hist. 6:1-549. DuRRANT, S. D., AND N. K. Dean. 1959. Mammals of Glen Canyon. Anthro. Papers, Univ. Utah 40:73-106. Hayward, C. L., D. E. Beck, and W. W. Tanner. 1958. Zoology of the Upper Colorado River Basin, I. The biotic communities. Brigham Young Univ. Sci. Bull., Biol. Ser. 1(3): 1-74. Hayward, C. L., and M. L. Killpack. 1958. Distribu- tion and variation of the Utah poulation (sic) of the Great Basin pocket mouse. Great Basin Nat. 18:26-30. Johnson, D. W. 1976. Populations of small mammals on isolated buttes in Canyonlands National Park, Utah. Unpublished thesis, Univ. Wyoming, La- ramie, 84 pp. Kelson, K. R. 1951. Speciation in rodents of the Colo- rado River drainage. Biol. Ser., Univ. Utah, ll(.3):vii 4- 1-125. Lee, M. R. 1960. Montane mammals of southeastern Utah— with emphasis on the effects of past cli- mates upon occurrence and differentiation. U :i- published dissertation, Univ. Utah, iv -t- 199 pp. LoHMAN, S. W. 1974. The geologic story of Canyonlards National Park. Bull., U.S. Geol. Surv., 1327: ix -i- 1-126. MacMahon, J. A. 1976. Species and guild similarity of North American desert mammal faunas: a func- tional analysis of communities. Pp. 133-148. In: Evolution of desert biota (D. W. Goodall, ed.), Univ. Texas Press, Austin, 250 pp. SoKAL, R. R., AND P. H. A. Sneath. 1963. Principles of numerical taxonomy. W. H. Freeman and Co., San Francisco, xvi -I- 359 pp. Tanner, W. W. 1965. A comparative population study of small vertebrates in the uranium areas of the Upper Colorado River Basin, Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 7(1): 1-31. Wilson, L., M. E. Olsen, T. B. Hutchings, A. R. Southard, and A. J. Erickson. 1975. Soils of Utah. Utah State Univ. Agric. Exper. Sta. Bull. 492: viii -t- 1-94. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo, Utah 84602. 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Authors publishing in the Great Basin Naturalist Memoirs may be expected to contribute $35 per printed page in addition to the cost of the printed copies they purchase. No printed copies are furnished free of charge. A price list for reprints and an order form are sent with the galley proof to contributors. Reprint Schedule of the Great Basin Naturalist 2 pp. 4 pp. 6 pp. 8 pp. 10 pp. 12 pp. Each 100 copies $20 $24 $28 $32 $36 $40 additional 200 copies 28 32 36 40 44 48 2 pp. 300 copies 36 40 44 48 52 56 $4 Great Basin Naturalist Memoirs No. 1 The birds of Utah. By C. L. Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. No. 2 Intermountain biogeography: A symposium. By K. T. Harper, J. L. Reveal, et al. $15. TABLE OF CONTENTS Review of tularemia in Utah and the Great Basin. Harold E. Stark 103 Chemical composition of some important plants of southeastern Utah summer ranges related to mule deer reproduction. Jordan C. Pederson and K. T. Harper 122 Emergence data and artificial rearing media for an aspen bark beetle, Tnjpophlociis popiili (Coleoptera: Scolytidae). David \. Stewart, Gary M. Booth, Jerold L. Petty ' ' 129 New svnonvmv and new species of .American bark beetles (Coleoptera: Scolvtidae), Part'vi'll. Stephen L. Wood '. 133 Annual energy budgets for three common rodent species in the northern Great Ba- sin. R. Kent Schreiber 143 Preliminary survey of raptor species in the Manti Division, Manti-LaSal National Forest. Stephen G. Jones 155 Vegetation response to a luoisture gradient on an ephemeral stream in central Ari- zona. Deborah .\nn Bloss and Jack D. Brotherson 161 Ecological and community relationships of Eriogonum con/nibosuin (Polygonaceae) in the Uinta Basin, Utah. Jack D. Brotherson and Karen J. Brotherson 177 Variation in hemoglobin types in the deer mouse (Peromyscus maniciikitiis) along an altitudinal gradient. David Wasserman and Donald J. Nash 192 First record of Patapiiis spinosii.s in Idaho and Nevada (Hemiptera: Leptopodidae). Donald R. Brothers 195 Bee visitation of Plilox hiyoiclcs (Polemoniaceae). V. J. Tepedino 197 Ecological distribution of rodents in Canyonlands National Park, Utah. David M. Armstrong 199 IHE GREAT BASIN NATURAUS illume 39 No. 3 September 30, 1979 FEB^ Brigham Young Univers '^^t'^.%T.-°u ^RY mo f^K '%l^'^ >C.l. X \ ■ %.|f>^ Xsi imuMMUMua GREAT BASIN NATURALIST Editor. Stephen L. Wood, Department of Zoology, Brigham Young University, Provo, Utah 84602. Editorial Board. Kimball T. Harper, Botany; Wilmer W. Tanner, Life Science Museum; Stanley L. Welsh, Botany; Clayton M. White, Zoology. Ex Officio Editorial Board Members. A. Lester Allen, Dean, College of Biological and Agricul- tural Sciences; Ernest L. Olson, Director, Brigham Young University Press, University Editor. The Great Basin Naturalist was founded in 1939 by Vasco M. Tanner. It has been published from one to four times a year since then by Brigham Young University, Provo, Utah. In gener- al, only previously unpublished manuscripts of less than 100 printed pages in length and per- taining to the biological and natural history of western North America are accepted. The Great Basin Naturalist Memoirs was established in 1976 for scholarly works in biological natu- ral history longer than can be accommodated in the parent publication. The Memoirs appears irregularly and bears no geographical restriction in subject matter. Manuscripts are subject to the approval of the editor. Subscriptions. The annual subscription to the Great Basin Naturalist is $12 (outside the United States $13). The price for single numbers is $4 each. All back numbers are in print and are available for sale. All matters pertaining to the purchase of subscriptions and back num- bers should be directed to Brigham Young University, Life Science 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. All manuscripts and other copy for either the Great Basin Naturalist or the Great Basin Naturalist Memoirs should be addressed to the editor as instructed on the back 11-79 650 42594 The Great Basin Naturalist Published at Provo, Utah, by Brigham Young University ISSN 0017-3614 Volume 39 September 30, 1979 No. 3 VEGETATIVE AND EDAPHIC FACTORS AFFECTING ABUNDANCE AND DISTRIBUTION OF SMALL MAMMALS IN SOUTHEAST OREGON George A. Feldhamer' Abstract.— The relationships between vegetative and edaphic habitat factors and the local distribution and abun- dance of small mammals on Malheur National Wildlife Refuge, Harney County, Oregon, were examined between July 1973 and June 1975. Of 16 species of small mammals captured, deer mice {Peromyscus maniculatus), montane voles {Microttts montanus). Great Basin pocket mice {Perognathus parvus), and least chipmunks {Eutamias minimus) comprised 90.1 percent of the individuals. The physiognomy of the vegetation was a factor in the distribution of rodent species other than deer mice. Pocket mice and chipmunks were restricted to the communities dominated by sagebrush (Artemisia tridentata) or greasewood (Sarcobatus vermiculatus). Population densities of pocket mice and chipmunks were significantly related to edaphic factors such as soil depth, texture, and strength, which may have affected the construction and stability of burrows. Montane voles occurred only in marsh or grassland communities. Population densities of voles were directly correlated with the amount of cover and inversely correlated with its patchiness. Deer mice were the most common species encountered and occurred in all but the grassland commu- nities. The density of this species was related to vegetative or edaphic factors only seasonally or in certain habitats, and few generalizations could be made. The general habitat preferences of many species of small mammals have been docu- mented by several generations of natural his- torians, and often can be attributed to food preferences and associated morphological ad- aptations (Baker 1971). Many other extrinsic factors affect populations of small mammals, including vegetation, soils, predation, com- petition, and weather. Intrinsic factors, such as genetic and behavioral changes, also are of significance (Krebs 1964:63-67). However, the quantitative relationships between many environmental factors and the distribution and abundance of small mammals usually are considerably less well known than their gen- eral preferences. This study was initiated to provide quan- titative information concerning the effect of vegetative and edaphic factors on the local distribution and abundance of small mam- mals inhabiting four community types on the Malheur National Wildlife Refuge. Description of the Area Malheur National Wildlife Refuge is pri- marily a resting and breeding area for migra- tory waterfowl and is located in the Harney Basin, Harney County, Oregon, between 118.5° and 119.5° W longitude and 42.7° and 43.4° N. latitude (Fig. 1). The refuge is at an elevation of approximately 1250 meters (m). The climate is characterized by dry sum- mers with temperatures rarely exceeding 32.2 C and cold winters with average temper- atures of -6.6 C. The average annual precipi- tation is 22.9 cm, much of which occurs as snowfall (Meteorology Committee, Pacific Northwest River Basin Commission 1969). Much of the refuge consists of valley wet- 'Department of Fisheries and Wildlife, Oregon State University, Corvallis, Oregon 97331. Present address: Appalachian Environmental Laboratory, Cen- ter for Environmental and Estuarine Studies, University of Maryland, Frostburg State College Campus, Gunter Hall, Frostburg, Maryland 21532. 207 208 Great Basin Naturalist Vol. 39, No. 3 lands vegetated primarily by hardstem bul- rush {Scirpus acuttis), cattail {Typha latifolia), baltic rush {Juncus balticus), sedges {Carex sp.), and submerged and emergent wetland flora. Rimrock areas above the valley floor support mainly big sagebrush, greasewood, and grasses, principally the exotic annual, cheatgrass (Bromus tectorum). Grassland areas were dominated by bluegrass {Poa se- cunda), saltgrass {Distichlis stricta), bluestem {Agropyron sniithii), and numerous other gra- minoid species, as well as sedges. Methods and Materials Eighteen study plots were established among the four predominant types of terres- trial plant communities on the refuge. Five plots each were in sagebrush and greasewood areas and four plots each were in marsh and grassland commimities. Areas designated as marsh were characterized by habitat factors that generally conformed to the "inland shal- low fresh marsh" category of Shaw and Fred- ine (1956:21). Areas designated as grassland conformed to the "inland fresh meadow" cat- egory. Live-trapping on all plots was con- ducted once during 1973 (July-September), twice in 1974 (June-August, Septem- ber-November), and once in 1975 (April-Jime). Each trapping period roughly corresponded seasonally to either spring, summer, or fall, except in 1973. Vegetative and edaphic parameters were measured dur- ing each trapping period. Fig. 1. Location of Malheur National Wildlife Ref- uge, Harney County, Oregon. Small Mammals Trapping grids consisted of 49 Sherman live-traps at 15.0 m intervals in a square grid of 1.1 ha (2.7 acres). This area included a bor- der of 0.3 ha (0.7 acres), from which it was assumed animals would be captured (Faust et al. 1971). Traps were operated for 10 consecutive days, except during 1973, when grids were operated for 3- or 4-day periods. Traps were baited with rolled oats, contained Dacron batting for nesting material, and were cov- ered with aluminum shields to minimize heat stress (Feldhamer 1977). Animals were re- moved from traps as soon after dawn as pos- sible. Trapped animals were individually marked by toe clipping, following the procedure out- lined by Taber and Cowan (1971), and re- leased at their respective points of capture. The species, sex, reproductive condition, age class, weight, and trap locality of each cap- tured animal were recorded. Reproductively active males had descend- ed testes. Females were termed pregnant if their abdomens were visibly swollen or if de- veloping young were detected by palpation. Nursing females were recognized by lacking fur around the teats or by large and protrud- ing mammae. Individuals were classified as juvenile or adult on the basis of body size and weight. All deer mice {Peromyscus manicukitus) with grey pelage or an incomplete postjuvenile developmental molt were considered juve- niles (Layne 1968). The estimated mean den- sities during each trapping period, and asso- ciated estimates of variance, were calculated using a Mean Peterson Estimate (Seber 1973:138). The trap-revealed distribution of the four common species inhabiting the study plots were classified as uniform, random, or clumped by South wood's (1966:36) index of dispersion (X'). Vegetation Percentage cover was estimated on each plot using a Gossen Tri-Lux photoelectric cell. Light intensity was measured between 1100 and 1300 hours at ground level and im- mediately above the vegetation at each of 10 Sept. 1979 Feldhamer: Oregon Mammals 209 points. The percentage differences between readings represented the amount of cover at each point. Measurements were made at 20 m intervals along two diagonal transects on each plot and the average considered an in- dex to the cover present during that trapping period. In addition, in 1975 each plot was visually "divided" into 49 quadrats, each 15.0 m- with a trap-site at the center. The per- centage of cover in each quadrat was esti- mated and rated as follows: 0-20 percent = 1; 21-40 percent = 2; 41-60 percent = 3; 61-80 percent = 4; and 81-100 percent = 5 (Myton 1974). For each species of small mammals, a chi-square ratio test was used to determine if the number of captures were equal for each of the five ratings. Expected values were calculated as— (number of total captures sites with E (captures/ "" P^"* rating i) rating i) trap sites These distributions were considered separate- ly for each plot. Although an average cover value was cal- culated for each plot, individual portions of- ten differed substantially in the amount of cover present. Therefore, the same five-divi- sion rating system (Myton 1974) was used to calculate a cover diversity index ("patch- iness") from the 10 photometric cover read- ings made on each plot. The formula l/2(f was used (M'Closkey and Fieldwick 1975) where q was the proportion of readings with- in each of the five cover rankings. During the initial trapping period on each plot in 1974, the foliage height diversity (FHD) of the vegetation was measured using general methods described previously (Mac- Arthur and MacArthur 1961, Rosenzweig and Winakur 1969, M'Closkey and Lajoie 1975). Vegetative density was measured in a differ- ent direction from each of the eight central trap-stations at heights of 7.6, 15.0, 30.0, 46.0, and 61.0 cm above the ground (q). The FHD was computed using l/Sq? with i = 1, 3, and 5 only. Thus, the vegetation was con- sidered to occupy three distinct layers: below 15.0 cm, between 15.0 and 46.0 cm, and above 46.0 cm. The percentage of vegetative moisture (succulence) was determined each trapping period beginning in 1974. Ten samples were collected at 20 m intervals along two diago- nal transects and an average succulence value per plot computed. The procedure involved clipping vegetation from an approximate 1 -m- area, placing samples in airtight cans, and weighing each sample before and after oven drying at about 63 C to determine the weight of water in the material. The ratio of water weight to dry weight was considered the percent moisture content (Dawson 1972). Soils The soil texture of each study plot was de- termined once in 1973 and was considered to remain constant for the duration of the field work. Ten soil subsamples were collected from depths of approximately 15.0 cm along diagonal transects. Subsamples were com- bined to form a single sample that was ana- lyzed for composition of sand, silt, and clay using the Bouyoucos method (Dawson 1971). The mean soil depth on each plot was esti- mated by forcing a sharpened steel probe, 1.0 cm in diameter, into the ground to a depth of 102.0 cm. If an obstruction was met prior to this depth, the distance from ground surface to the obstruction was recorded. An average depth was determined from 10 probings made along diagonal transects. The soil sheer stress ("strength") on each plot was measured once during the initial trapping period of 1974, using a Soil Test pocket penetrometer. The index value, mea- sured in kg/cm-, was in direct proportion to the soil strength. Ten measurements, made along diagonal transects, were averaged to obtain a mean value. Diversity values for both soil depth and soil strength were calcu- lated using l/Sqf. For soil depth, diversity was computed on the basis of four equal rankings (q) of 25.4 cm each. Diversity for soil strength also was computed on the basis of four rankings: 0-1.12, 1.12-2.25, 2.26-3.39, and 3.40-4.50 kg/cm^ The percentage soil moisture on each plot was estimated each trapping period follow- ing the procedure described to estimate vege- tative succulence. Ten samples were collect- ed from a depth of 15.0 cm along diagonal transects and an average value computed. 210 Great Basin Naturalist Vol. 39, No. 3 Regression Analyses Linear regression analyses were used to de- termine the relationship between the esti- mated mean density of each species of small mammal and concurrently measured inde- pendent habitat parameters. Habitat varia- bles and associated densities of small mam- mal species in each of the four community types were analyzed in three groupings: (1) for the entire study period; (2) for all periods exclusive of 1973, so that the effects of vege- tative succulence and patchiness could be evaluated; and (3) for the initial trapping pe- riod of 1974, because foliage height diversity and soil strength were measured only during this period. Correlations were considered to be statistically significant if ?< 0.05. malian orders (Table 1). Three orders were represented by single species and were con- sidered incidental to the study because traps were not set for them. Although 13 species of rodents from four families were captured, deer mice, montane voles {Microtus mon- tanus). Great Basin pocket mice (Perognathus parvus), and least chipmunks {Eutamias min- i7nus) comprised 90.1 percent of the small mammals captured and were considered in detail in the analyses and discussion. All four species had trap-revealed sex ratios signifi- cantly biased toward males (P<0.05). Male- to-female ratios were: deer mice, 1.7:1.00; montane voles, 2.12:1.00; Great Basin pocket mice, 1.7:1.00; and least chipmunks, 1.9:1.00. Perognathus parvus Results and Discussion During the field work, 26,460 trap-nights on the study plots produced 4,717 captures of small mammals. These captures involved 1,580 individuals and represented four mam- Density.— Great Basin pocket mice were resident only in sagebrush or greasewood communities. In sagebrush communities, among-plot variation in the density of pocket mice differed seasonally only by a factor of about 4. Within-plot variation did not exceed Table L Total number of individuals of each mammalian species captured on study plots in the four predominant types of terrestrial plant communities on Malheur National Wildlife Refuge from July 1973 through June 1975. Species Community Type Order Sagebrush^ Greasewood'' Marsh^ Grassland'^ Total Rodentia Peramyscus maniculatus 153 231 (245f 59 (86) 1 (2) 444 Microtus montanus 0 7 (7) 330 (478) 100 (171) 437 Perognathus parvus 206 72 (76) 5 (7) 0 283 Eutamias minimus 101 159 (169) 0 0 260 Reithrodontomys megalotis 1 13 (14) 36 (52) 21 (36) 71 Dipodomys ordii 30 11 (12) 0 0 41 Dipodomys microps 9 2 (2) 0 0 11 Onychomys leucogaster 7 0 0 0 7 Microtus longicaudus 0 0 3 (4) 0 3 Microdipodops megacephalus 3 0 0 0 3 Spermophilus townsendii 2 1 (1) 0 0 3 Neotoma lepida 2 0 0 0 2 Thomomys talpoides 1 0 0 0 1 Carnivora Mustela frenata 0 1 (1) 2 (3) 0 3 Insectivora Sorex vagrans 0 0 10 (14) 0 10 Lagomorpha Sylvilagus nuttallii TOTALS 1 516 0 0 0 1 497 (528) 445 (645) 122 (209) 1,580 hko 0.605 0.549 0.389 0.207 ^Involved 5 plots and 8,232 trap nights. "Involved 5 plots and 7,742 trap nights. '^Involved 4 plots and 5,684 trap nights. "Involved 4 plots and 4,802 trap nights. ^Because unequal effort was expended in each habitat type, numbers and grassland habitats, based on a total effort comparable to that made in 'Calculated as H' = l/N(logjonl-21ogjQnjl) according to Lloyd et al. 1! n parenthesis represent relative totals for animals captured in greasewood, marsh, sagebnish areas. 168, for the rodent species in each community type. Sept. 1979 Felohamer: Oregon Mammals 211 a factor of 5 seasonally. In the greasewood areas, however, among-plot differences in the density of pocket mice differed by as much as ;i factor of about 7; there was an e(juivalent difference in density within plots (Table 2). Although densities were generally lower on the greasewood than the sagebrush plots (t = 4.29, df = 37, F<0.001), in both habitat types peak numbers of pocket mice were trapped from late April to early June. The lowest population densities occurred from late June through mid-August. These fluctua- tions in population density were similar in both timing and magnitude to those of a pop- ulation of P. parvus studied by O'Farrell et al. (1975) in shrub-steppe habitat in southeast Washington. Densities of pocket mice appar- ently increased on most plots in the fall. This was probably the result of increased activity as temperatures cooled. Reproduction.— Male pocket mice were in breeding condition from early May until early August. Peak breeding activity prob- ably occurred in early June, a period not ade- quately represented by trapping data. Males were believed to be reproductively active about a month before the females, based on the percentage in reproductive condition in May (56 percent of males, 8 percent of fe- males). Pregnant females were trapped al- most exclusively in June, although sample size was small (n = 4), and no reproductively active pocket mice were trapped after 29 August in either habitat type. Juvenile pocket mice were trapped from early May to early September on plots in sagebnish areas, although the majority were found from June through August. In grease- wood areas, juveniles were trapped only from June through August (Fig. 2). Recruitment of juveniles was greatest in both habitat types during July. Dispersion.— The trap-revealed dispersion of pocket mice on all plots differed signifi- cantly from random, and a clumped distribu- tion pattern was evident. On each of three plots where 60 or more capture records of pocket mice were obtained, a significant relationship was evident between pocket mouse distribution and cover density. Signifi- cantly fewer pocket mice than expected were " trapped in quadrats with less than 40 percent cover, but more pocket mice than expected were trapped in quadrats with greater than 40 percent cover (X- = 26.33, df = 8, P<0.001). The preference of pocket mice for areas of relatively dense cover undoubtedly was a re- sponse to the interaction of several factors. Increased vegetation possibly reduced the rate of predation (Rosenzweig and Winakur 1969, Brown and Lieberman 1973) and in- creased forage availability (O'Farrell 1975). It also probably beneficially affected micro- habitat evaporation rates, humidity, and air temperature (Beatley 1976). Habitat.— Although vegetation was of ob- vious importance in the local distribution of pocket mice, the abundance of this species in 100 80 60 40 20 0 100 80 60- 40- 20 Greasewood 5i2 i i Sagebrush 38^1 2515 6i2 17:11 5^ 5^ i I May Jun JuT Aug Sep Oct Nov Fig. 2. Percentage of captures of pocket mice each month comprised of adult (open rectangles) and juvenile (solid rectangles) animals. Numbers atop open rectangles denote maleiemale sex ratios. 212 Great Basin Naturalist Vol. 39, No. 3 all sagebrush or greasewood areas was signifi- cantly correlated only with edaphic factors. A direct correlation between the population density of pocket mice and the percentage of sand on each plot was evident (P< 0.05— Fig. 3A). The ability of pocket mice to dig through the surface layer of the soil is of ob- vious importance to their fossorial activities. The general importance of edaphic factors was further suggested by an inverse relation- ship between density and the percentage of clay in the soil (P<0.05-Fig. 3B). These fac- tors probably affect burrow construction and stability. For example, soil texture has a di- rect influence on several aspects of soil mois- ture, including depth and rate of percolation, retention, and evaporation rates (Krynine 1947, Beatley 1976). In greasewood areas the mean soil moisture was inversely correlated with percentage of sand (r- = 0.92, P<0.005). However, Rosenzweig and Winakur (1969) found the distribution and ab mi dance of five other species of pocket mice in Arizona showed no correlation with soil texture. Eiitamias minimus Density.— Least chipmunks also occurred only in sagebrush or greasewood commu- nities. In the sagebrush community type, among-plot differences in the density of chip- mimks generally were less than a factor of 3. Within-plot fluctuations in population den- sity varied by an equivalent amount. In greasewood, among-plot variation in popu- lation density varied seasonally by a factor of about 6, with the exception of the 1973 trapping period. There was an equivalent variation within-plot seasonally (Table 2). In neither shrub community type was there a season during which peak numbers were evi- dent. Vaughan (1974) also noted a fairly stable population density for this species in northern Colorado. Reproduction.— Male chipmunks may have been reproductively active in both commu- nity types about one month before the fe- males, as suggested by the percentage of each sex in breeding condition in May (91 percent of males, 40 percent of females). Breeding apparently was confined to a fairly brief pe- riod, with peak activity in late April and May. The number of chipmunks in breeding condition declined rapidly in both commu- nity types throughout the summer. No repro- ductively active chipmunks were trapped af- ter 31 July in the greasewood areas or after 29 August in sagebrush areas. Davis (1939) and Gordon (1943) reported that this species mated from early to midspring in the north- ern part of its range, and Negus and Findley (1959) reported no sexually active least chip- munks occurring after late June in northwest Wyoming. Of the 260 chipmunks captured during the study, only one was considered a juvenile. Juveniles were probably most numerous in early June, when no trapping was conducted. This is suggested by the reproductive data Pb 10 20 30 40 50 60 70 80 PERCENTAGE SAND IN SHRUB PLOTS 10 20 30 40 50 60 70 80 PERCENTAGE CLAY IN SHRUB PLOTS Fig. 3. Relationship between components of soil tex- ture and mean density of pocket mice on 10 study plots in shnib communities on Malheur National Wildlife Rcf uge from July 1973 through June 1975. (A) Percentage "t sand; (B) Percentage of clay. Sept. 1979 Feldhamer: Ore{;on Mammals 213 above and tlie re.sults of Hall (1946) and Lins- dale (1938), who reported parturition in least chipmunks in Nevada occurred during May and early June. Tevis (1958) found gravid least chipmunks during mid-April in north- eastern California. It is po.ssihle the age cri- teria may have been inadequate to di.s- tinguish juvenile chipmunks. Dispersion.— On all plots where more than 23 capture records of chipmunks were ob- tained, their dispersion was clumped. On none of the plots was there a relationship be- tween dispersion and the amount of vegeta- tion cover. However, on only one plot were there as many as 60 location records. Habitat.— On both .shrub areas, the density of chipmunks was directly correlated with the mean depth of soil (P<0.05, Fig. 4) and with soil strength (P<0.05, Fig. 5). On sage- bmsh areas, a positive correlation was found between density and diversity of soil strength (r- = 0.88, P<0.01). Correlations between the density of chipmunks and habitat factors on the greasewood areas included a direct relationship to the percentage of clay in the soil (r- = 0.72, P<0.05). As with pocket mice, the depth, texture, and strength of the soil would directly affect chipmunks in the construction and stability of burrows, and indirectly affect aspects of temperature and humidity. The densities of pocket mice and chipmunks were oppositely related to the percentage of clay in the soil; this suggests that chipmunks did not find it as difficult as the smaller pocket mice to dig through a sometimes hard, con.solidated soil surface of high clay fraction. Peromyscus maniculatus Density.— Deer mice exhibited the widest local distribution, being resident in sage- brush, greasewood, and marsh community types (Table 1). Deer mice generally were most abundant on the greasewood areas, where among-plot variation differed by a fac- Table 2. Estimated population densitie.s and standard deviation for four species of small mammals common on studv plots (1.1 ha) in three community tvpes on Malheur National Wildlife Refuge from July 1973 through June 1975. Plot Sageb rush Greasewood Marsh P. man. P. pa, rvus E. min. P. man. P. pai rvus E. min. P. man. M. mont. Period N i : Sd N ± Sd N i : Sd N ± : Sd N ± Sd N ± Sd N ± Sd N ± Sd 1973 1 4.8 0.4 5.5 0.9 7.7 3.2 9.3 2.0 8.7 2.7 1.7 0.7 5.6 0.9 0.0 - Julv- 2 13.7 2.4 5.5 0.5 2.3 0.3 9.2 1.9 0.0 _ 16.9 1.7 b b Sept. 3 2.3 0.3 8.4 0.4 2.0 0.6 6.2 1.2 0.7 0.7 6.5 2.6 0.0 - 169.5 .30.0 4 1.0 0.0 9.0 2.0 0.0 — 2.0 I.O 0.0 _ 14.3 0.3 0.0 252.8 82.2 5 3.2 0.2 9.5 1.5 0.0 - 5.3 0.3 2.0 - 0.0 - - - 1974 1 2.8 0.4 9.3 0.9 7.9 1.1 6.5 0.9 8.3 0.9 2.1 0.7 5.2 1.0 0.0 - June- 2 6.9 0.5 12.5 1.4 9.7 1.6 10.1 1.8 0.0 — 9.8 0.9 0.0 - 0.0 - Aug. 3 0.8 0.3 5.8 0.8 3.6 0.4 9.1 0.7 1.3 0.2 10.7 1.6 4.7 0.8 46.6 5.4 4 0.8 0.1 6.0 0.5 0.0 — 4.4 0.4 0.0 _ 9.8 0.5 1.3 0.6 .36.5 4.7 5 2.9 0.1 4.4 0.7 0.0 - 4.4 0.5 4.0 0.3 0.0 - - - - 1974 1 6.3 0.5 15.1 2.6 6.4 0.6 7.2 0.4 11.9 0.8 5.1 0.8 8.2 1.1 0.0 - Sept.- 2 5.7 0.3 11.6 1.6 9.4 0.7 20.1 1.7 0.0 _ 13.6 2.3 0.0 _ 0.1 - Nov. 3 3.0 0.0 5.4 0.2 5.9 0.7 17.0 1.0 1.8 0.2 6.6 0.8 6.4 0.5 33.9 6.7 4 8.5 0.6 5.0 0.2 0.0 _ 7.8 0.9 1.0 _ 11.6 1.2 5.4 1.6 40.2 13.9 5 24.4 0.4 6.7 0.3 0.0 - 4.3 0.9 0.7 - 0.0 - - - - 197.5 1 0.8 0.1 10.8 0.9 13.9 1.0 3.5 0.5 3.4 0.3 5.9 0.7 a a April- 2 2.9 0.3 16.3 0.6 6.0 0.6 a a a 0.0 0.0 June 3 3.0 0.3 4.8 0.2 2.2 0.4 18.2 1.1 8.7 0.3 4.4 0.5 1.4 0.2 4.2 0.8 4 14.8 0.8 12.0 0.8 0.0 — 15.0 0.9 3.9 0.8 3.5 0.4 a a 5 11.8 0.9 12.3 0.3 0.0 - 20.5 1.9 10.7 0.7 0.0 - - - - *Plot was flooded, could not be operated. ''Grid not operated during 197.3. 214 Great Basin Naturalist Vol. 39, No. 3 tor greater than 3 only in 1973 (Table 2). Within-plot fluctuation in the density of deer mice also approached this magnitude. Popu- lations of deer mice in sagebrush areas exhib- ited fluctuations in density comparable to those on greasewood areas. Both within- and among-plot variation generally differed by a factor of about 3. The densities of deer mice on sagebrush and marsh plots were about equal, with fluctuations in the marsh areas again relatively minor. There was no season during which the density of this species was Mean density chipmunks (y) = -1.278 « 0095 soil depth (x) .2 10 20 30 40 50 60 70 80 90 100 110 SOIL DEPTH (cm) Fig. 4. Relationship between the density of chipmunks and soil depth on study plots in two shrub habitat types on Malheur National Wildlife Refuge from July 1973 through June 1975. Mean density chipmunks (y } - -2.352 * 6.347 soil strength (x) tec )4 TSeoS'^'^TrTTT^nT'^l^TS^S SOIL "STRENGTH" (kg/cm') Fig. 5. Relationship between the density of chipmunks and soil "strength" on study plots in two shnib habitat types on Malheur National Wildlife Refuge from July 1973 through June 1975. Sept. 1979 Feldhamer: Oregon Mammals 215 consistently highest, although density was generally lowest during the summer. Reproduction.— Deer mice were reproduc- tively active during all months that trapping was conducted, although a decline in breed- ing activity was evident during the summer. Parturition occurred at least as early as April in the shrub habitats, because juvenile ani- mals were trapped in May (Fig. 6). Peak numbers of juveniles were on the plots in May and in the fall. It was not determined if breeding continued throughout the year, but, considering the usually harsh winter condi- tions on the study area, it seems doubtful. However, deer mice in sagebrush areas of east-central Washington have bred through- out the year (Scheffer 1924). Dispersion.— The trap-revealed dispersion of deer mice in all habitat types differed sig- nificantly from random, and on 10 of 11 plots a clumped pattern was evident. Relationships between dispersion and cover in the marsh plots could not be evaluated because of too few captures in 1975. The results of this anal- ysis on sagebrush and greasewood plots were equivocal. On greasewood plot 4 and sage- brush plot 5, significantly fewer deer mice than expected were captured in portions of those plots with less than 40 percent cover, but greater numbers than expected occurred in portions with more than 40 percent cover (X^ = 58.59, df = 8, P< 0.001). However, on sagebrush plot 2 the opposite relationship was apparent (X- = 7.53, df=l, P<0.01), and on the remaining five plots, where sufficient capture records were available to allow anal- ysis, there were no significant relationships between the amount of cover and the dis- persion of deer mice. These results reflect the range of relation- ships relative to cover previously reported for this species. A direct relationship between amount of cover and the local distribution and abundance of Peromyscus was described by Allred and Beck (1963). However, the pro- portion of cover did not exceed 25.0 percent in any of the areas sampled by these authors. An inverse relationship between cover and density was reported for several habitat types, including grassland and cultivated areas (Phillips 1936, LoBue and Darnell 1959, Tester and Marshall 1961), desert shrub (MacMillen 1964), and sites disturbed by strip-mining (Dusek and McCann 1975). In similar habitats, other researchers found no relationship between amount of cover and lo- cal distribution or abundance of deer mice (Rickard 1960, Verts 1957). It may be that no general relationship exists between the local distribution or abundance of deer mice and amount of cover. Habitat.— There was a direct correlation between the density of deer mice and only one vegetative factor: the amount of vegeta- tion at the three heights used to measure fo- liage height diversity. This correlation was evident on plots in both sagebrush areas (r- = 0.98, P<0.01) and greasewood areas (r^ = 0.86, P<0.25). Thus, at least during the 100 80 60- 40 ^ 20 c 0) ° 100 S) 80 o 40 201- 0 Greasewood i i i 815 i 21 Sagebrush 107 I I 1 May Jun Jul Aug Sep Oct Nov Fig. 6. Percentage of captures of deer mice each month comprised of adult (open rectangles) and juve- niles (solid rectangles) animals. Numbers atop open rec- tangles denote male:female sex ratios. 216 Great Basin Naturalist Vol. 39, No. 3 summer, it appeared that deer mice in shrub communities selected for increased foliage, at least at the measured, interspersed levels of the vegetative profile, rather than a contin- uum of foliage above or below a particular height. Whether this relationship was of di- rect adaptive significance in predator avoid- ance, foraging or general scansorial tenden- cies (Horner 1954), or was indirectly associated with some other factor, was not readily apparent. In marsh areas, a slight positive correlation existed between the density of deer mice and the percentage of sand in the soil (r- = 0.37, F<0.05). On greasewood areas, an inverse relationship was evident between these two factors (r^ = 0.80, F<0.025), as well as a di- rect correlation between density of deer mice and the percentage of soil moisture (r- = 0.30, P<0.01). Therefore, deer mice exhibited an opposite response to the percentage of sand on marsh and greasewood areas, and no rela- tionship on sagebrush areas; habitats general- ly subjected to large differences in the amount of free water associated with them. The permeability and drainage capabilities of soils vary directly with their percentage of sand (Krynine 1947). The opposite response of deer mice to the percentage of sand on marsh and greasewood plots may have re- sulted from an effort by deer mice to select an "optimal" moisture range within the con- tinuum of soil moisture conditions that could be tolerated. That is, "wetter" arid areas and "drier" wet areas. Microtus montanus Density — In marsh areas, montane voles were resident only on plots 3 and 4, which were dominated by burreed {Sparganium sp.), as opposed to bulrush {Scirpus paludosis) and spikerush {Eleocharis palustris) on plots 1 and 2. The density of voles during 1973 was the highest of any small mammal during the study. Densities declined sharply after the in- itial trapping period. In 1975, the population' density of voles on marsh plot 3 was reduced from the 1973 estimate by a factor of at least 20 (Table 2). Although the population den- sities of voles in grassland areas were general- ly lower than in marsh areas, the grassland plots were continuously altered by land-use practices throughout the field work, and trends in the density of voles were difficult to determine. Reproduction.— The reproductive data for voles were not as complete as those for other species of small mammals. Reproductively active voles were trapped in marsh areas only from July through September. However, montane voles were imdoubtedly breeding during the spring months (Bailey 1936), when trapping was not conducted in marsh or grassland communities. Also, considering the short gestation period of this species (Asdell 1964), and that juvenile voles were trapped in early November, breeding must also have occurred in October. There was no period during which the juvenile increment of the population or the percentage of adults in breeding condition was consistently largest. Dispersion.— The trap-revealed dispersion of voles on all plots differed significantly from random, and a clumped pattern was evident. On the only plot where the analysis could be made, the dispersion of voles was related to the amount of cover. Voles were trapped significantly more often in quadrats where cover was greater than 80 percent (X^= 15.49, df=l,F<0.005). Habitat.- There was a weak, direct corre- lation between the estimated population den- sities of voles and the mean amount of cover on plots in marsh (r- = 0.41, P<0.01) and grassland communities (r- = 0.41, P<0.025). In a related manner, an inverse relationship was evident between the density of voles and the patchiness of cover of these habitat types (r- = 0.41, P<0.01 and r^ = 0.64, P O % 13 ^21 'A '/^ Vy^ Collecting Date y^ % 15 - Fig. 5. Niche overlap indices for Ceratium hirundinella and Aphanizomenon flos-aquae during the summer of 1974. 252 Great Basin Naturalist Vol. 39, No. 3 Literature Cited Cole, G. A. 1975. Textbook of limnology. C. V. Mosby Co., St. Louis. 283 p. CoLWELL, R. K., AND D. J. FuTUYMA. 1971. On the mea- surement of niche breadth and overlap. Ecology 52:567-576. Darwin, C. 1859. The origin of species. John Murray, London. 513 p. Cause, G. F. 1969. The struggle for existence. Hafner Publ. Co., New York. Originally publ. 1934. 163 P- George, D. G., and R. W. Edwards. 1973. Daphnia dis- tribution within Langmuir circulations. Limnol. and Oceanogr. 18:798-800. Hardin, G. 1960. The competitive exclusion principle. Science 131:1292-1297. Harvey, H. W. 1969. Chemistry and fertility of sea- waters. Cambridge Univ. Press, Cambridge. 240 P- Horne, a. J., and R. C. Wrigley. 1975. The use of re- mote sensing to detect how wind influences planktonic blue-green algal distribution. Verb. Internat. Verein. Limnol. 19:784-791. Hutchinson, G. E. 1961. The paradox of the plankton. Am. Nat. 95:137-145. Jaeger, R. G. 1971. Competitive exclusion as a factor in- fluencing the distribution of two species of terres- trial Salamanders. Ecology 52:632-637. Klecka, W. R. 1975. Discriminant analysis. Pages 434-467. In: H. H. Nie, C. H. Hull, J. G. Jenkins, K. Steinbrenner, D. H. Brent. SPSS Statistical package for the social sciences. McGraw-Hill Co., New York. Levandowsky, M. 1972. Ecological niches of sympatric phytoplankton species. Am. Nat. 106:71-78. MacArthur, R. H. 1958. Population ecology of some warblers or north-eastern coniferous forests. Ecol- ogy 39:599-619. Makarewicz, J. C, and G. E. Likens. 1975. Science 190:1000-1003. Miracle, M. R. 1974. Niche structure in freshwater zoo- plankton: a principle components approach. Ecology 55:1306-1316. Petersen, R. 1975. The paradox of the plankton: an equilibrium hypothesis. Am. Nat. 109:35-49. RiCHERSON, P., R. Armstrong, and C. R. Goldman. 1970. Contemporaneous disequilibrium, a new hypothesis to explain the paradox of the plank- ton. Proc. Nat'l. Acad. Sci. 67:1710-1714. ScHOENER, T. W. 1968. The Anolis lizards of Bimini: re- source partitioning in a complex fauna. Ecology 49:704-706. Snedecor, G. W., and W. G. Cochran. 1967. Statistical methods. Iowa State Univ. Press, Ames. 593 p. Stumm, W., and J. J. Morgan. 1970. Aquatic chemistry. Wiley-interscience, New York. 583 p. Titman, D. 1976. Ecological competition betwen algae: experimental confirmation of resource-based competition theory. Science 192:463-465. Whiting, M. C, J. D. Brotherson, and S. R. RusHFORTH. 1978. Environmental interaction in summer algal communities of Utah Lake. Great Basin Nat. 38:31-41. GROUND NESTING AND AGGRESSIVE BEHAVIOR BY THE SWAINSON'S HAWK {BUTEO SWAINSONI) Neil D. VVoffiiiden^'2 and James A. Mosher'-' .\bstract.— a successful ground nesting by Swainson's Hawks, Biiteo sivainsoni, is reported for central Utah. Un- usual aggressive nest defense behavior is described for this ground nesting pair. The Swainson's Hawk (Biiteo swainsoni) is a large Buteo of western North America, nesting ahiiost exclusively in trees (Bent 1937, Life Histories of North American birds of prey, part 1, U.S. Natl. Mus. Bull. No. 167). In central Utah, on 9 May 1973, we found a Swainson's Hawk nest that was con- structed on a rock ledge approximately 1.5 m above the surrounding ground. The ledge sloped gently to the ground providing easy access to the nest. The nearest tree was ap- proximately 700 m to the southwest. The nest was composed primarily of dry sticks, both woody and herbaceous, with green, leaf-bear- ing branches from juniper {Juniperiis os- teospertna) and atriplex {Atriplex spp.). The cup was lined with strips of juniper bark and grass. Three young were successfully fledged from this ground nest. The nest was used only during the 1973 nesting season. In both 1974 and 1975 Swain- son's Hawks successfully nested in the small juniper tree, mentioned above. The tree nest was constructed by refurbishing a Ferru- ginous Hawk {Buteo regalis) nest that had previously been built in the 2 m tree. It is not known if the same pair of hawks were in- volved in all three nestings; the premise ap- pears likely, however, based on the proximity of the two nest sites. Two young hawks fledged from the tree nest in 1974; one fledged in 1975. It would appear that, be- cause of their greater vulnerability, ground nests would be less productive than tree nests. In this particular case, we feel that any increased vulnerability was compensated by the additional aggressiveness of the ground nesting hawks. However, it should be pointed out that ground nesting Ferruginous Hawks appear to be as reproductively successful as tree nesting pairs. Bent (1937:231-232) characterized the Swainson's Hawk as "a gentle, unobtrusive bird, living in harmony with its feathered neighbors both large and small." He further stated that this species is not aggressive and "has rarely been known even to threaten to attack an intruder" at the nest. Dunkle (1977, Auk 94:65-71) found Swainson's Hawks nest- ing in Wyoming to vary in their aggressive- ness toward humans, with some females com- ing to within a few meters of the intruder while others stayed many meters away. Our nest visits to this Great Basin ground nest elicited highly aggressive behavior from both members of the pair. Attacks were in the form of long, shallow dives that brought the hawks to within a few centimeters of us. These were frequent and intense, and contin- ued until we retreated 30 to 40 m from the nest. On one occasion, while we were banding the three young, the adults failed to initially react with the usual degree of intensity. A few weak passes were made, after which the female soared to a great height and contin- ued to soar in a tight circle overhead. The male flew from view behind a hill. The fe- male then began a vertical descent, falling rapidly for perhaps 100 m. Pulling out of the 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 'Current address; Division of Natural Sciences, University of Pittsburgh at Johnstown, Johnstown, Pennsylvania 15904. 'Current address; Appalachian Environmental Laboratory, GEES, University of Maryland, Frostburg, Maryland 21532. 253 254 Great Basin Xatur-\list Vol. 39, No. dive, she circled for a few seconds and began a second descent. At this time the male flew in along the face of the rock outcrop adjacent to the nest ledge, striking the investigator at the nest in the back of the head. Tmning 90 degrees, he continued his attack, forcing the second investigator to the grovmd. He then joined the female who had terminated her dive and was circling above us. They then drifted slowly away from the nest site. One interpretation of this observation is that nest defense behavior may be a plastic trait and the degree of aggressiveness is re- lated to the security of the nest site. We would encourage others to be alert to this possible relationship between nest defense behavior and nest site security. We wish to thank J. Edward Gates for re- viewing this manuscript. This is Contribution No. 592 Appalachian Environmental Labora- torv' Center for Environmental and Estuarine Studies, University of Mar\land, Frostburg State College Campus, Gunter Hall. Frost- burg, Maryland. RESPONSE OF REPTILE POPULATIONS TO DIFFERENT LAND MANAGEMENT PRACTICES ON THE IDAHO NATIONAL ENGINEERING LABORATORY SITE Timothy D. Reynolds' Abstract.— Populations of reptiles were examined in grazed and ungrazed habitats dominated by sagebrush {Arte- misia tridentata) or by crested wheatgrass (Agropyron cirstatum) on the Idaho National Engineering Laboratory (INEL) Site in southeastern Idaho. The sagebnish lizard (Scelopoms graciosus) and the short-horned lizard {Phryno- soma douglassi) were the only species of reptiles encountered in sufficient numbers to permit statistical analysis. Both of these species preferred sagebrush habitats to areas dominated by crested wheatgrass. The sagebrush lizard was most abundant in the native, ungrazed, sagebrush habitat, and the short -horned lizard was most plentiful in the sheep-grazed area dominated by big sagebrush. Sagebrush, mostly big sagebnish {Artemisia tridentata), was once the dominant vegeta- tion over 100 milHon ha of western range- lands in the United States (Beetle 1960). In this century, at least 10 percent of the sage- brush range has been altered in an attempt to increase forage production for livestock (Braim et al. 1976). In Idaho, approximately 850 thousand ha of public and private range- land, most of which was dominated by big sagebrush, have been chemically or mechani- cally treated and reseeded with crested wheatgrass (Agropyron cristatum and A. desertonan) (BLM document 1974, L. Sharp, pers. comm.). With few exceptions, studies concerning the effects of the alteration of sagebrush communities on wildlife have concentrated on game species such as mule deer {Odo- coileus hemionns), pronghorn antelope {An- tilocapra americana), elk {Cervus canadensis), and Sage Grouse (Centrocercus urophasianus) (Patterson 1952, Mackie 1965, Anderson 1969, Peteron 1971). The effects of altering the sagebrush habitat on nongame species have received little attention (Braun et al. 1976). Reptiles, especially lizards, are important components of North American ecosystems, rivaling mammals and birds with respect to numbers, biomass, and energetics in the desert community (Turner et al. 1976). This vertebrate group is frequently overlooked in environmental studies. The objective of this study was to evaluate the response of reptile populations to different land management practices on the Idaho National Engineering Laboratory (INEL) Site. Study Sites This study was conducted on the INEL National Environmental Research Park Site in southeastern Idaho. The INEL Site oc- cupies 2,315 km- of cool desert on the Snake River Plain, approximately 48 km west of Idaho Falls, Idaho (Fig. 1). The topography of the site is flat to gently rolling, with fre- quent lava outcroppings characteristic of the Columbia Plateau Province (Atwood 1970). The elevation ranges from 1454 to 1554 m, with the median about 1500 m. The climate is characterized by hot summers and cold winters, with record high and low temper- atures of 39 and -41 C. The mean annual temperature at the Central Facilities Area (CFA) (Fig. 1) is 5.8 C. Precipitation, mostly in the form of spring rains, averages 21.6 cm annually, with a 30-year range of 11.4-36.3 cm. The vegetation of the INEL Site is domi- nated by big sagebrush (Jeppson and Holte 1976). Four areas of different land use were se- lected for study. Two of these were domi- 'Department of Biology, Idaho State University, Pocatello, Idaho 8; Operations Office, 550 South Second Street, Idaho Falls, Idaho 83401. Present Address: Environmental Sciences Branch, Department of Energy, Idaho 255 256 Great Basin Naturalist Vol. 39, No. cm IDAHO 28 'DAHO 22 KILOMETERS F ■ '■ ■ I 0 8 PAVED ROAD [ I UNPROTECTED [ I PROTECTED IDAHO FALLS Fig. 1. Location of the grazed and ungrazed study areas on the INEL Site used in determining the response of | reptile populations to different land management practices. The grazed and ungrazed sites each contained a habitat dominated by sagebrush {Artemisia tridentata) (SAGE) and crested wheatgrass (Agwpywn cristattim) (CWG). Those portions of the INEL Site protected from live-stock grazing are shown. Sept. 1979 Reynolds: Idaho Reptiles 257 nated by big sagebmsh, and two were former sagebrush range reseeded with crested wheat- grass. One sagebrush and one crested wheat- grass study area were 1.3 km northwest of Atomic City, Idaho, just within the southern boundary of the INEL Site (Fig. 1). The crested wheatgrass here was planted in 1958 or 1959 (INEL Site records vary) after the native vegetation was removed by disking. These two areas were contiguous and have been grazed by 1000-1200 domestic sheep each spring since 1960. These study areas are referred to as grazed sage and grazed CWG. The other sagebrush and crested wheat- grass study areas were approximately 12 km northwest of the grazed areas (Fig. 1). The sagebrush in this area (ungrazed sage) had re- ceived no grazing pressures since 1950, and served as the control plot for this study. The adjacent crested wheatgrass planting (un- grazed CWG) was seeded in 1960, after win- ter flooding inundated and killed the native vegetation (A. Olson, pers. comm.). Livestock were last permitted in this area 10 years pri- or to the planting of crested wheatgrass. Materials and Methods Vegetation was sampled in each study area along 400 m of line transects with 20 X 50 cm sampling frames (Daubenmire 1959). The percent coverage for each plant species was estimated using six cover classes (0-5, 5-25, 25-50, 50-75, 75-95, and 95-100). The mid- points of each class were used to calculate the average canopy coverage for each spe- cies. These values were used in calculating the percent similarity (PS) of the vegetal communities among the study areas (Orloci 1967). The invertebrate fauna was sampled in an attempt to determine the potential prey available to reptiles in each area of different land use. A 4-ha-square grid was established in each of the study areas, and sweep netting was performed along four transect lines par- allel to each axis of the grids. Sweep samples were taken using a standard 38 cm (15 in) canvas sweep net. All transects were swept twice, resulting in 5600 sweeps in each study area. Pit-fall (sink) traps and a "noose on a stick" (Linder and Fichter 1977) were used to cap- ture lizards. The sink traps were number 10 tin cans buried in the ground at 50 m inter- vals in a 4-ha grid system, for a total of 25 pitfall traps in each study area. Masonite cov- ers (23 X 23 X 0.5 cm) were placed over the traps when not in use. Initially, these covers were removed for periods not exceeding 4 hours, thus precluding heat prostration of any reptiles in the trap. I found that trapping suc- cess was improved and trap mortality re- duced to zero when the lids were not totally removed, but propped open slightly with a stick. Thus, the traps were effectively open continuously, but the lids shaded any animals in the trap. Traps were checked at 1- or 2- day intervals from May through October 1977. The noose consisted of a small copper wire or monofilament fishing line fashioned into a slipping noose and attached to the tip portion of a fishing rod. All lizards captured by either method were color-marked with in- delible ink for individual identification. Serpentine species, with the exception of rattlesnakes (Crotalus viridis), were hand-cap- tured and color-marked when they were en- countered. When a rattlesnake was found, its size was estimated and recorded along with the location of the sighting and any dis- tinguishing characteristics of the snake. The paucity of snakes encountered, and the small number of lizards recaptured, prohibited meaningful estimates of population size by conventional methods. Therefore, the num- ber of reptiles encountered was taken as an indication of the relative density of each spe- cies in each of the habitats studied. Results and Discussion As expected, big sagebrush was the domi- nant plant species in the sage study areas, and crested wheatgrass was the dominant species on both CWG areas (Table 1). The ungrazed sage had a significantly greater vegetal species diversity (H = 2.85) than did any other area (Table 2). The two CWG study areas exhibited a greater floristic per- cent similarity (PS = 90 percent) than any other habitat comparisons (Table 2). Similar numbers of invertebrates were col- lected in the ungrazed and grazed habitats dominated by sagebrush (181 and 185, re- spectively) (Table 3). Equal sweep-netting ef- 258 Great Basin Naturalist Vol. 39, No. forts resulted in the capture of approximately 50 percent more invertebrates in each of the crested wheatgrass stands than in either of the sagebrush areas. These differences were significant (X- 05(i) = 90.2). Four species of reptiles were encountered during the study (Table 4). No serpentine species were found in either of the ungrazed study areas. Gopher snakes {Pituophis me- lanoleucus) and rattlesnakes are generally common and widespread over the INEL Site (Linder and Sehman 1977). The absence of snakes in the imgrazed study areas may not be an accurate indication of the effects of dif- ferent land use practices on these species. Woodbury et al. (1951), Hirth et al. (1969), and Sehman (1977) recorded maximum dis- persal distances of snakes from a hibernacu- lum of 2.0, 1.3, and 1.9 km, respectively. It is likely that the ungrazed study areas were Table 1. Percent coverage (%) and frequency (f) of plant species in grazed and ungrazed habitats dominated by sagebrush {Artemisia tridentata) and by crested wheatgrass (Agropyron cristatum) on the INEL Site. Only those spe- cies with a frequency of 10 or more, or a percent coverage of at least 1 percent are included. Ungrazed Grazed Crested C rested Sagebrush wheatgrass Sa |ebrush wh eat grass % f % f f % f Opuntia polyacantha 1.3 13 Chenopodium leptophyllum 0.6 17 Artemisia tridentata 17.00 105 25.00 113 0.5 5 ■■ Aster scopulorum 0.7 23 Chrysothamnus nauseosus 0.2 3 1.6 15 0.4 5 Arabis spersiflora 0.3 11 1.0 15 Agropyron cristatum 1.9 15 52.0 160 39.4 154 Agropyron spicatum 6.0 35 Oryzopsis hymenoides 5.2 34 0.2 3 Poa sanbergii 1.8 16 Sitanion hystrix 3.0 35 9.2 93 Astragalis purshii 1.9 16 Phlox hoodii 1.4 11 Habgeton glomeratus 1.4 4 3.9 13 Total number of species 31 3 6 5 Total percent coverage 41.6 53.2 38.7 44.1 Species diversity (H) 2.85 0.37 1.17 0.54 Table 2. Percent similarity (PS) and species diversity (H) comparisons of vegetation in grazed and ungrazed habitats dominated by sagebrush (Artemisia tridentata) and by crested wheatgrass (Agropyron cristatum) on the INEL Site. Numbers in the upper right portion of the body of the table are PS determinations. Those in the lower left are the results of t-test analysis of species diversity. (H) Percent similarity U ngrazed Grazed Diversity Sagebrush Crested wheatgrass Sagebrush Crested wheatgrass Ungrazed Sagebrush Crested wheatgrass Grazed Sagebrush Crested wheatgrass 2.85 0..37 1.17 0.54 6.78° 20.78° 8.72° 33 2.18° 0.47 38 41 2.37° 32 90 22 •H significantly different as P <0.05. Sept. 1979 Reynolds: Idaho Reptiles 259 more than 2 km from a suitable denning site. Therefore, snakes were exchided from further analysis. Two species of lizards were frequently en- countered: the short-horned lizard {Phryno- wina douglassi) and the sagebrush lizard Scehporus graciosus). The relative density of 3oth species was significantly greater in each )f the sagebrush study areas than in either .rested wheatgrass area (Table 5). Signifi- L'antly more (X-05(i) = 6.92) sagebrush lizards ^ere encountered in the ungrazed sage than n the grazed sage (Table 5). Conversely, sig- lificantly more (X-o5(i) = 4.76) short-horned lizards were found in the grazed sage than in its ungrazed counterpart. The two crested wheatgrass study areas supported populations of both species of lizards that were not statis- tically different (Table 5). When the data from the ungrazed sage and ungrazed CWG study areas were combined and compared with the combined data from the two grazed study areas, sagebrush lizards were found to occur significantly more often (X".o5(i) = ^-00) in the absence of grazing. Phrynosoma douglassi population levels were not statistically different (X^o5(i)==2.67) when analyzed in the same manner. Table 3. Number of invertebrates collected in grazed and ungrazed habitas dominated by sagebrush {Artemisia tri- dentata) and crested wheatgrass {Agrapyron cristatum) on the INEL Site. Taxon Ungrazed Grazed Sagebrush Crested wheatgrass Sagebrush Crested wheatgrass Arachnoidea Unclassified HOMOPTERA Ceropidae Ciradellidae Aphidae Other Unclassified Hemiptera Corizidae Miridae Other Unclassified COLEOPTERA Chrysomelidae Other Unclassified Orthoptera Acrididae Other 3 119 7 8 44 7 40 19 15 0 5 0 2 1 0 2 0 0 10 0 6 0 3 1 6 0 12 8 5 22 1 15 2 1 2 1 0 0 0 12 2 1 5 10 11 10 12 0 4 13 1 6 3 3 6 2 Lepidoptera Unclassified Diptera Other Unclassified Hymenoptera Formicidae Other Unclassified Total 0 0 1 2 18 25 11 51 27 24 26 23 0 0 0 7 19 37 36 53 181 270 185 276 260 Great Basin Naturalist Vol. 39, No. 3 Conclusions Populations of lizards responded differen- tly to different land management practices on the INEL Site. Grazing by sheep in a hab- itat dominated by sagebrush resulted in a sig- nificant reduction in the plant diversity of that area. This was not accompanied by a concommitant change in the relative density of either the invertebrate or reptilian fauna. Planting a former sagebrush range with crested wheatgrass resulted in a further re- Table 4. Number of reptiles encountered in grazed and ungrazed habitats dominated by sagebnish (Artemisia tri- dentata) and by crested wheatgrass (Argopyron cristatum) on the INEL Site. Number encountered Ungrazed Grazed Taxon Sagebrush Crested wheatgras; Sagebrush Crested wheatgrass Short-homed hzard (Phyronosoma douglassi) Sagebrush hzard (Sceloporus graciosus) Gopher snake {Pituophis melanoleucus) Great Basin rattlesnake (Crotalus viridis) 37 Table 5. Results of chi square analysis of the number (N) of sagebnish lizards (Sceloporus graciosus) and short- horned lizards (Phrynosoma douglassi) encountered in grazed and ungrazed habitats dominated by sagebrush (Arte- misia tridentata) and by crested wheatgrass (Argopyron cristatum) on the INEL Site. Study area Ungrazed Grazed Species and number encountered Sagebrush Crested wheatgrass Sagebnish Crested wheatgrass Short -horned lizard (Phryonosoma douglassi) Ungrazed Sagebrush Crested wheatgrass Grazed Sagebrush Crested wheatgrass (17) (4) (26) (7) 9.94' 4.76* 16.13' 5.88' 0.82 10.94° Sagebnish lizard (Sceloporus graciosus) Ungrazed Sagebrush Crested wheatgrass Grazed Sagebrush Crested wheatgrass (37) (5) (21) (0) 27.68' 9.58' 36.01° 3.20 19.05° •Significant at P < 0.05. Sept. 1979 Reynolds: Idaho Reptiles 261 ){ duction in the diversity of vegetation. This a was accompanied by an apparent increase in V the relative density of invertebrate fauna (e.s- i pecially arachnids, dipterans, and hynien- li opterans), although the populations of lizards • were significantly below that found in the sagebrush habitats. The increase in in- vertebrates in the crested wheatgrass plan- ,i tings may indeed be a very real phenomena, jr it may indicate a shortcoming in using ■iweep-nets as a sampling technique in habi- tats that are structurally different. Ants are the primary prey items of both the sagebrush lizard (Burkholder and Tanner 1974) and the short-horned lizard (Guyer 1978), and it is interesting to note that formi- L'ids were in equal abundance in each of the four habitats studied (Table .3). Thus, if prey ilone was the factor controlling the distribu- :ion of lizards, one would expect equal den- sities of lizards in each of the four habitats of different land use. This, of course, was not the case, indicating that the physiognomy of the vegetation, as well as the presence of po- - tential prey, is vital in determining the den- sity of saurians, and most likely reflects the coevolution of these lizard species with the sagebrush environment. Grazing a sagebrush habitat appeared to improve that habitat for P. douglassi, where- - as S. graciosus population levels seemed to be lowered by livestock grazing. The mecha- nism(s) responsible for this disparity in the re- sponse of these species to grazing practices is not known. It is possible that the few forbs and grasses in the grazed sage study area (Table 1) resulted in more, or improved, bas- king sites for the short-homed lizard, but the lack of ground cover had a negative effect on the sagebrush lizard populations. The precise microhabitat affinities of these lizards are not recorded in the literature. Thus, the results presented here may reflect abiotic and biotic differences, as yet imdetermined, among the four habitats studied. In any case, converting a sagebrush range to crested wheatgrass re- sulted in the reduction of the population lev- els of both the short-horned and sagebrush Uzards. And, once the native vegetation was replaced by crested wheatgrass, grazing did not appear to further affect the lizard popu- lations. As the production of agricultural products increases to keep pace with the ever increas- ing demand for red meat protein, it is likely that more and more acreage of habitat domi- nated by sagebrush will be converted to mon- ocultures of crested wheatgrass. Additionally, efforts to reclaim strip-mined lands in the In- termountain West rely heavily on the plan- ting of crested wheatgrass for initial soil stabilization. Thus, crested wheatgrass plan- tings are becoming more widespread in our environment. This study, and others like it (Reynolds 1978), indicate that the continued conversion of habitats dominated by sage- bmsh to large expanses of crested wheatgrass will have a long-term and deleterious effect on the populations of native vertebrates. Acknowledgments I thank C. Guyer, M. Mahoney, and M. Reynolds for assistance in installing and regu- larly monitoring the sink traps. S. Condie and J. Jensen provided invaluable assistance in the collection and classification of the in- vertebrate fauna. Drs. J. E. Anderson, E. Fichter, O. D. Markham, and C. H. Trost critically evaluated an earlier draft of the manuscript. J. and G. Millard offered useful suggestions. This is a contribution from the INEL Site Radioecology-Ecology Studies Program, administered by the U.S. Depart- ment of Energy. Literature Cited Anderson, A. E. 1969. 2,4-D, sagebrush, and mule deer- cattle use of upper winter range. Colorado Divi- sion of Game, Fish and Parks, Special Report No. 21. 21 p. Atwood, N. D. 1970. Flora of the National Reactor Testing Station. Brigham Young Univ. Sci. Bull., Biol. Ser. 11(4): 1-46. Beetle, A. A. 1960. A study of sagebrush: the section Tridentatae of Artemisia. Univ. Wyoming Agric. Expt. Sta. Bull. .368, Laramie. 83 p. Braun, C. E., M. F. Baker, R. L. Eng, J. S. Gashwiler, AND M. H. Schroeder. 1976. Conservation com- mittee report on effects of alteration of sagebrush communities of the associated avifauna. Wilson Bull. 88:165-171. Bureau of Land Management. 1974. Idaho range man- agement program report, p. I-6-I-7. Burkholder, G. L., and W. W. Tanner. 1974. Life his- tory and ecology of the Great Basin sagebrush swift, Sceloporus graciosus graciosus Baird and Girard, 1852. Brigham Young Univ. Sci. Bull., Biol. Ser. 19(5): 1-44. 262 Great Basin Naturalist Vol. 39, No. 3 Daubenmire, R. F. 1959. A canopy-coverage method of vegetational analysis. Northwest Sci. 33:4.3-64. GuYER, C. 1978. Comparative ecology of the short- horned lizard (Phrijnosoma douglassi) and the sagebmsh lizard {Scelopowus gmciosus). Unpub- lished thesis, Idaho State Univ. 130 p. HiRTH, H. F., R. C. Pendleton, A. C. King, and T. R. DowNARD. 1969. Dispersal of snakes from a hi- bernaculum in northwestern Utah. Ecology 50:332-339. Jeppson, R. J., and K. E. Holte. 1976. Long-term vege- tation analysis of the INEL Site. Pages 160-165. In: 1975 Progress Report Idaho National Engi- neering National Site Radioecology-Ecology Pro- grams. IDO-12080. Idaho Operations Office U.S. ERDA. LiNDER, A. D., and E. Fighter. 1977. The amphibians and reptiles of Idaho. Idaho State Univ. Press, Pocatello, Id. 78 p. LiNDER, A. D., AND R. W. Sehman. 1977. Herpetofauna of the Idaho National Engineering Laboratory Site. J. Idaho .Acad. Sci. 13:42-46. Mackie, R. J. 1965. Range ecology and relations of mule deer, elk and cattle in the Missouri River Breaks, Montana. Unpublished dissertation. Montana State Univ. Orloci, L. 1967. An agglomerative method for classifi- cation of plant communities. J. Ecol. .55:19.3-205. Patterson, R. L. 1952. The sage grouse in Wyoming. Wyoming Game and Fish Comm., and Sage Books, Inc., Denver, Colorado. 341 p. Peterson, M. M. 1971. From sagebrush to waving grass. Soil Cons. .36:210-211. Rey.nolds, T. D. 1978. The response of native vertebrate populations to different land management prac- tices on the Idaho National Engineering Labora- tory Site. Unpublished dissertation. Idaho State Univ. 105 p. Sehma.n, R. W. 1977. Hibernaculum dynamics of the Great Basin rattlesnake {Crotaltis viridis lutosus). Unpublished thesis. Idaho State Univ. 66 p. Turner, F. B., P. A. Medica, and B. W. Kowalewsky. 1976. Energy utilization by a desert lizard {Ufa stansburiana). US/IBP Desert Biome Monograph No. 1. Utah State Univ. Press, Logan. Woodbury, A. M., B. Vetas, G. Julian, H. R. Glissmyer, R. L. Heyrend, A. Call, E. W. S.mart, and R. T. Sa.nders. 1951. Symposium: A snake den in Tooele County, Utah. Herpetologica 7:1-52. DECREASES OF JUNIPER WOODLAND IN THE UTAH AND SALT LAKE VALLEYS SINCE SETTLEMENT E. M. Christensen' and J. D. Brotherson- .\bstract.— Although pinyon-juniper woodland is common throughout most of Utah, it is less so in Utah and Salt Lake valleys. Even so, stands do occur in the valleys and are known to have been there since presettlement times. Three such stands occurring along the eastern sections of the valleys have shown major decreases in area since settle- ment times, and one has all but disappeared. Their disappearance is attributed to early use for fuel, agricultural, and urbanization purposes. The expansion since pioneer days of pin- yon-juniper woodland in the foothills and benchlands of Utah is a well-known and documented phenomenon. Parker (1971) and Barney (1972) summarized the literature con- cerning the invasion of pinyon-juniper wood- land that was earlier described by Mason (1963) as taking place on deep soils, relative- ly free of stones, and which formerly "sup- ported a dense cover of grass, forbs, and shrubs." In the eastern part of the Salt Lake and Utah valleys, however, an opposite trend has occurred: a decrease of junipers and juniper woodland. Discussed here are decreases in three es- tablished stands of juniper as described by early pioneer residents. Pinyon pine {Piniis edulis Englem.) does not occur in Salt Lake Valley and is rare in Utah Valley (Erdman 1970). Therefore, the woodlands here de- scribed are essentially composed of Utah jimipers {Jtiniperus osteosperma (Torr. Little), also known locally as "cedar." Part of Salt Lake City's folklore indicates that there was only one tree in the valley, a Utah juniper located at 300 South and 600 East Streets (Young 1930, Carter 1940), al- though other trees were noted by Brigham Young in 1847 (Young 1930). The stump of this particular tree stood for many years after the tree died in 1933 and part of it is still preserved under a monument canopy. Juni- pers were apparently not abundant in the Salt Lake Valley at the time of settlement be- cause most early descriptions found in diaries and reports do not mention them (Wakefield 1933, 1936). There were some junipers scat- tered throughout the valley, however, as is shown in the writings of Langworthy (1855), Bryant (1949), and Gunnison (1852). Lang- worthy observed Salt Lake Valley from the southeast on 11 July 1850. He was impressed by the scarcity of trees, but noted the con- ifers on the hillsides: . . . The whole expanse looks bleak and naked, there being no trees in sight except a very few along the banks of streams, and some stunted, scattering pines and ce- dars on the sides of the mountains. Historical Distribution of Juniper It is generally believed that at the time of settlement, the Salt Lake Valley had few na- tive trees on the foothills and benchlands (Smith 1953, Roberts 1965), although trees are known to have been common along the streams (Wakefield 1933, Wakefield 1936). Bryant, traveling through the Salt Lake Valley in 1849, also recorded the occurrence of junipers on the sides of the mountains: . . . Hidden away in the profound chasms and along the streams whose beds are deeply worn in the moun- tain sides are the cedar, pine, dwarf-maple, and occa- sionally oak, where the inhabitants of the vale seek their fuel and building timber ... 'Dr. E. M. Christensen, Professor of Botany and Range Science, Department of Botany and Range Science, Brigham Young University, had gathered the material for this paper before his death in 1973. It is here pubHshed posthumously. ^Department of Botany and Range Science, Brigham Young University, Provo, Utah 84602. 263 264 Great Basin Naturalist Vol. 39, No. 3 The situation in Utah Valley was much the same as that in Salt Lake Valley. Cottam, the first ecologist to describe the vegetation in Utah Valley, commented that on the "west- ern escarpment of the Wasatch . . . except for white fir near the summit, the absence of trees is noticeable" (Cottom 1926). Carvalho, who saw Utah Valley some 70 years earlier on 6 May 1854 (Carvalho 1860) stated: . . . The scenery, which is enUvened by . . . flowering vales ... is without timber except on the creeks which meander from the mountains . . . Sparse growths of young Cottonwood are the only trees I have seen except in the cannons of the mountains, on which grow pines, cedars, and a species of mahogany. Huntington (1842), an early pioneer, de- scribed Utah Valley in 1849 in his diary: . . . Besides the Salt Lake Valley are several smaller valleys, the largest of which is the Ewtah, which would be the end of my searches to find a pleasant and delight- ful home . . . The most of the western side of the valley is . . . covered with juniper trees . . . the borders of . . . creeks and rivers are well stocked with wood. There are jimiper stands with old trees on the hillsides near the mouths of several can- yons in both valleys: Parley's Canyon, Mill Creek Canyon, Tolcat's Canyon, Big Cotton- wood Canyon, Spanish Fork Canyon. These groves are rather open and the older trees are usually highlined from browsing. There is little reproduction in the stands, but it ap- pears sufficient to maintain the present den- sity of the existing stands. These juniper stands were doubtless present in pioneer days, and are the types of stands referred to in the descriptions given above. Decrease of Presettlement Juniper Stands Point of the Mountain Stand.— There were several well-established juniper stands of various sizes in Salt Lake and Utah valleys when settlement began. One of these was at Point of the Mountain near Jordan Narrows on the Traverse Range (T4S, RIW, S 13, 23, 24). The stand was generally ignored in the descriptions of the early travelers and pio- neers, probably because the road bypassed it. This stand was still present in 1933 when Wakefield studied the ecology of early Salt Lake and Utah valleys (Wakefield 1933), but Wakefield did not note its size. Prior to the development of the gravel pits and the con struction of Interstate Highway 1-15 during | the past 15 years, there was an open grove of juniper trees remaining on the hill east of the road. Presently, there are only about 20 to 30 trees, which are scattered along the tops of the road cut of the 1-15 Highway and the ad- jacent gravel pits to the north. Another remnant stand of junipers also ex- ists a few miles northeast of Lehi at Cedar Hollow, located about half a mile east of Lehi Cemetery (T5S, RIE, S3,4). The history of the community is unknown to the authors, but there are a few juniper trees there at the present time. Mapleton Stand.— In 1926 Cottam de- scribed only one stand of junipers in eastern Utah Valley (which is an indication that it was the largest one in the valley). He ob- served: . . . the juniper belongs largely to the mountains. It is found sparingly on Mapleton Bench, but it reaches asso- ciation rank on the slopes of Lake and West mountains. By "association rank" Cottam meant that the species was a dominant of a major plant community. Lake and West mountains are on the western boundary of Utah Lake. By 1933 Wakefield noted "but a few remnant areas [of juniper communities] on Mapleton Bench." Observations of early pioneers give the im- pression that in presettlement times this juni- per stand was more extensive than either the brief descriptions of Cottam or Wakefield in- dicate. Mae B. Huntington summarized the impressions of the first pioneers of the Springville area with considerable emphasis on vegetation (Huff 1947): When the Pioneers alighted from their wagons that September afternoon in 1850, one pioneer mother said it looked like a heaven on earth. The weary travelers stood knee-deep in native grass. They saw the benchlands nearby covered with cedars. They saw the distant hills covered with cottonwood and balsam. In a history of Springville, Johnson (1900) relates an incident of the O. B. Huntington party in February 1849, when the horses strayed one night from a campsite in Spring- viUe toward the mouth of Maple Canyon "through the cedars which grew on what is now known as Mapleton Bench." Much of Springville and Mapleton occupy the original area that supported junipers, but Sept. 1979 Christensen, Brotherson: Juniper Decline 265 today only a remnant of the grove (mostly south of Evergreen Cemetery, T8S, R3E, S9) remains to remind us of the stand that the pioneers knew. Manila Stand.— This stand is located about 1.5 miles north of Manila and about a mile east of the Alpine Country Club (T4S, R2E, S31; T5S, R2E, S6). It is restricted to a west-facing slope (approximately 25 degrees) south of American Fork Canyon and west of Highway 146. Although it was not mentioned by Cottam or Wakefield, the stand was evi- dently present when settlement occurred. A ring count on one medium-sized tree (2.8 dm. diameter at 1 foot above the surface) in- dicated that the tree was over 190 years old in 1963. The stand was apparently rather large at one time. On 15 December 1850, George A. Smith was in a grove of junipers located between American Fork and Provo, where he "found about 50 wagons camped in the cedars." He may have been referring to the Manila Stand. If so, the grove must have been larger than it is today to have accom- modated 50 wagons, and it must have extend- ed onto the surrounding level ground. If Smith was in another stand of junipers, it has completely disappeared because no other groves of junipers exist between the two cities today. In either case, a decrease of juni- per woodland is indicated. In 1849, a group of 50 men led by Parley P. Pratt may have camped in this same grove of junipers. Colton (1946) gave the account of Robert T. Campbell, the clerk of the com- pany who wrote on 25 November 1849: The company traveled to Cedar Grove where they camped at 4 or 5 p.m. about two and one-half miles from the Utah Fort. Two days later they reached Provo River. It is about 12 miles from the Manila Stand to the site of Utah Fort and about 9 miles to the mouth of Provo Canyon, so either Campbell's estimation of distance was poor, the grove was much larger than today, or the group was in another "Cedar Grove" nearer to Pro- vo that no longer exists. In 1963 E. M. Christensen studied the Ma- nila Grove. It was then about one mile long and one-quarter of a mile wide. Today it cov- ers less than one-third of the area it covered in 1963, having been reduced primarily by gravel mining, road building, and a tract housing development. In 1963, there were 90 juniper trees per hectare (stems 1 dm in di- ameter at 1 foot above surface and larger) and 21 juniper plants per hectare smaller than tree size. The projected foliar cover of the ground surface of the junipers was 11 percent. The average diameter of the trees was 2.4 dm (9.4 inches at 1 foot above the ground. The stand has been heavily grazed, gully errosion is evident, and exotic species are common in the understory. With the present housing development under construc- tion, this stand will most likely disappear within the next 10 to 15 years. Summary A conclusion can be made that there were juniper woodland stands in the eastern sec- tions of Salt Lake and Utah valleys at the time of settlement by white men. Decrease in the area occupied by at least three of these juniper groves has occurred, even though in- creases in juniper have occurred elsewhere in Utah. In addition, juniper trees were scat- tered throughout these valleys, and, doubt- less, the number of such trees has also de- creased. Urbanization, agriculture, fuel procurement, etc., appear to be responsible for at least some of the decrease. Literature Cited Barney, M. A. 1972. Vegetation changes following fire in the pinyon-juniper type of west central Utah. Unpublished thesis, Brigham Young University, Provo, Utah. 71 p. Bryant, E. 1949. What I saw in California. D. Appleton & Company, New York. 490 p. Carter, K. B. (compiler). 1940. Trees, flowers and birds of Utah. Heart throbs of the West, Vol. 2. pp. 237-260. Daughters of the Utah Pioneers, Salt Lake City, Utah. 532 p. Carvalho, S. N. 1860. Incidents of travel and adventure in the far west. Derby and Jackson, New York. 380 p. Colton, R. C. 1946. A historical study of the explor- ation of Utah Valley and the story of Fort Utah. Unpublished thesis, Brigham Young University, Provo, Utah. 190 p. Cottam, W. P. 1926. An ecological study of the flora of Utah Lake, Utah. Unpublished dissertation. Uni- versity of Chicago, Chicago, Illinois. 46 p. Erdman, K. S. 1970. Distribution of the native trees of Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 11(3): 1-34. 266 Great Basin Naturalist Vol. 39, No. 3 Gunnison, Lieut. J. W. 1952. The Mormons, or Latter- day Saints, in the valley of the Great Salt Lake: A history of their rise and progress, peculiar doc- trines, present condition, and prospects, derived from personal observation, during residence among them. Lippincott, Grambo & Co. 168 p. Huff, E. N. (compiler). 1947. Memories that live: Utah County Centennial history. Daughters of the Utah Pioneers of Utah County. Art City Pub- lishing Co., Springville, Utah. 488 p. Huntington, O. B. 1942. Diary of Oliver B. Huntington, 1847-1900 Vol. 2. Typewritten copy in Brigham Young University Library. Provo, Utah. 455 p. Johnson, D. C. 1900. A brief history of Springville, Utah, from its first settlement September 18, 1850, to the 18th day of September 1900. Wil- liam F. Gibson, Springville, Utah. 124 p. Langworth, F. 1855. Scenery of the plains, mountains and mines. J. C. Sprague, Ogdensburgh, New York. 324 p. Mason, L. 1963. Using historical records to determine climax vegetation. J. Soil and Water Con- servation 18:190-194. Parker, R. D. 1971. A study of the effects of two con- version treatments on pinyon-juniper vegetation in Utah. Unpublished dissertation, Brigham Young University, Provo, Utah. 255 p. Robert, B. H. 1965. A comprehensive history of The Church of Jesus Christ of Latter-day Saints— Cen- tury I. Vol. III. Brigham Young University Press, Provo, Utah. 572 p. Smith, G. A. 1956. Journal of George Albert Smith (1817-1875). Typewritten copy in Brigham Young University Library, Provo, Utah. 86 p. Smith, J. F. 1953. Essentials in church history. 13th ed. Deseret Book Company, Salt Lake City, Utah. 722 p. Wakefield, J. H. 19.33. A study of the plant ecology of Salt Lake and Utah valleys before the Mormon immigration. Unpublished thesis, Brigham Young University, Provo, Utah. 57 p. . 1936. A study of the native vegetation of Salt Lake and Utah valleys as determined by histori- cal evidence. Proc. Utah Acad. Sci., Arts and Let- ters 13:11-16. Young, J. R. 1930. Early recollections of the Salt Lake Valley. Utah Historical Quart. 3:8.3-84. COMPETITION BETWEEN HARVESTER ANTS AND RODENTS IN THE COLD DESERT' Dan S. Landeeii^"^, Clive D. Jorgensen-^ and H. Diiane Sniith'^ Vbstract.— Local distribution patterns of three rodent species {Perognathus parvus, Perormjscus manictihitus, Rcithwdontomys megalotis) were studied in areas of high and low densities of harvester ants (Pogommiyrmex outjhcci) in Raft River Valley, Idaho. Numbers of rodents were greatest in areas of high ant-density during May, but partially reduced in August; whereas, the trend was reversed in areas of low ant-density. Seed abundance was prob- abl\ not the factor limiting changes in rodent populations, because seed densities of annual plants were always great- er in areas of high ant-density. Differences in seasonal population distributions of rodents between areas of high and low ant-densities were probably due to interactions of seed availability, rodent energetics, and predation. Competition for food is an important de- terminant of ecosystem structure and dynam- ics (Hairston et al. 1960, Brown and David- son 1977), but, because the relative importance of competition as a determinant of community composition is debatable (Schoener 1974, Wiens 1977), additional study is essential to provide necessary insights into community relationships. Most studies of competition have dealt with interactions be- tween species of closely related taxa (Connell 1961, Brown 1971, Sheppard 1971, Grant 1972, MacArthur 1972, Pianka 1974, Schro- der and Rosenzweig 1975), but only recently have they included species of more distantly related taxa (Hansen and Ueckert 1970, Brown et al. 1975, Fenton and Fleming 1975, Primack and Howe 1975, PuUiam and Brand 1975, Sinclair 1975, Brown and Davidson 1977, Boyden 1978). Although studies of harvester ants in North America have included distribution (Hull and Killough 1951, Sharp and Barr 1960, Cole 1968), damage and control (Cole 1932, Sever- in 1955, Lavigne 1966, Race 1966, Wight and Nicholes 1966), foraging activity (Cole 1934, Creighton 1953, Tevis 1958, Bernstein 1974, Rogers 1974, Whitford and Ettershank 1975, Whitford 1976), species diversity (Da- vidson 1977a, 1977b, Mares and Rosenzweig 1978), and taxonomy (Cole 1968), only a few studies in Arizona have examined com- petition between ants and rodents (Brown et al. 1975, Pulliam and Brand 1975, Brown and Davidson 1977, Reichman, unpubl. ms.). Reichman (unpubl. ms) demonstrated that ei- ther taxon alone or both together impact the density of seeds in the soil, but Pulliam and Brand (1975) concluded that little com- petition exists between ants and rodents ex- cept in years of low seed production. Brown and Davidson (1977) concluded that ants and rodents do compete for seeds in southern hot deserts, and harvester ants can affect rodent distribution. Our objective was to investigate local dis- tribution patterns of rodents and ants to de- termine if competition for seeds was an im- portant factor in any observed interaction between western harvester ants and rodents in the cold desert. Study Sites and Methods This study was conducted in a sagebrush- greasewood community at Raft River Valley, Idaho, from May through August 1977 and 1978. Predominant plant species were sage- brush {Artemisia tridentata), greasewood {Sar- cobatus vermiculatus), squirreltail barley (Si- tanion hystrix), and mustards (Lepidium perfoliatum and Descurainia richardsonii). Harvester ants were of the species Pogono- 'This research was sponsored by EG & G, Idaho, Inc., and supported by the U.S. Department of Energy Contract DE-A5O7-77ID0I674. ^Present address: Rockwell Hanford Operations, Richland, Washington 99352. 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 267 268 Great Basin Naturalist Vol. 39, No. 3 myrmex owyheei. Rodent species investigated were the white-footed deer mouse (Per- omysciis manicidatus), western harvest mouse (Reithrodontomys megalotis), and Great Basin pocket mouse {Perognathiis par- vus). One 12 X 12 base grid with 15 m between stations, and 10 4 X 5 mini-grids with 10 m between stations were estabUshed to assess ant-rodent interactions. Approximately half of the base grid and five of the mini-grid were located in areas of high ant-density, but the remainder of the base grid and the other five mini-grids supported only low ant-den- sities. Two Utah live traps (Garcia et al. 1974) baited with rolled oats were placed at each station on the base grid, which was trapped for 10 consecutive nights during May and August 1977 and 1978. The 10 mini-grids, with one Utah live trap at each station, were trapped simultaneously for four consecutive nights each month. Mini-grids were trapped prior to the base grid trapping in May and August. All animals on the base grid and mini-grids were marked for identification by clipping the toes and released after species, sex, and trap station were recorded. Numbers of rodents for the mini-grids and centers of activity on the base grid were estimated us- ing Hayne's (1949a, 1949b) methods. Understory vegetation (<30.5 cm high) and overstory vegetation (>30.5 cm high) were measured using the cover class and point quarter methods, respectively. Relative seed abundance and time of seed drop of an- nual forbes {Descurainia richardsonii, Lepi- dium perfoliatum) and grass {Sitanion hystrix) was determined monthly on the 10 mini-grids from soil samples, using the method of Franz et al. (1973). In order to establish a relationship between distribution of P. manicidatus and seed abun- dance during the May reproductive period, sex ratios were calculated for specimens col- lected from the 10 mini-grids. Ant mound volumes were used in con- junction with numbers of mounds per unit area to estimate ant density at each mini-grid (Fig. 1). Ant foraging activity was estimated twice each day during the first week of each month by placing a one-meter quadrat over each of 22 stations located 15 m apart in the area of high ant-density of the main grid. Numbers of active ants/m-/minute were counted and averaged to obtain a monthly es- timate of relative foraging activity. Seed preferences of harvester ants were deter- mined on 23 June and 6 and 28 July 1978 for five mounds. Seeds were collected, identified, and counted from 75 returning foragers for each mound at each sampling period. Results Areas of high ant-density had less total cover (32.8 percent and 42.2 percent in high and low ant-density areas, respectively) and less grass cover (0 and 32.7 percent in high and low ant-density areas, respectively) than did areas of low ant-density. Densities of seeds from annual plants {Descurainia rich- ardsonii, Lepidium perfoliatum) were appar- ently dropped in late May and early June, whereas Sitanion hystrix seeds were dropped in July and August (Table 1). This temporal difference in seed drop was also reflected by the foraging activities of the ants (Table 2). In June, ants almost exclusively foraged on seeds from D. richardsonii and L. per- foliatum, whereas in July they foraged on S. hystrix (Table 2). Ant foraging activity was lowest in May, peaked in July, and decreased in August (Table 3). There was a significant negative correlation (p< 0.025) between ant densities and rodent numbers (Fig. 2), but no signifi- cant correlation between rodent numbers and seed abundance (p< 0.200). Average numbers of rodents in areas of low ant-density were 9 ±3.2 rodents/ grid in May and 16 ±2.7 in August, an 88 percent in- crease; in areas of high ant-density the num- bers of rodents were 26 ±8.3 in May and 11 ±2.7 in August, a 58 percent decrease (Table 4). Centers of rodent activity on the | base grid shifted from the area of high ant- density to the area of low ant-density during August 1977 and 1978 (Table 4). Sex ratios of P. maniculatus during the May reproductive period on the five mini- grids with low seed-density averaged 1.6:1 (1:1 to 2:1) males to females, and on the five grids with high seed-density they averaged 1:1.3 (5:11 to 13:8). When fitted to a bino- mial distribution these ratios were signifi- Sept. 1979 Landeen et al.: Ant-Rodent Competition 269 cantly different (p<0.08) in both high and low seed-density areas. The female-to-male sex ratios were also significantly greater (p<0.04) in the areas of high seed-densities. Discussion Studies of competition by Brown and Da- vidson (1977) between ants and rodents in Arizona indicate that seeds limit the distribu- tion of either taxon, which also influences each other. Although they demonstrated sig- nificant evidence for exploitation com- petition between ants and rodents, our data from Idaho show that factors other than seed abimdance are more influential in changing rodent distributions in areas of high ant-den- sity. Our data indicate that D. richardsonii and L. perfoliatiwi seeds were always more abundant in areas of high ant-densities than they were in areas of low ant-densities (Table 1). Because this condition persisted in spite of greater foraging pressure by ants and rodents, these seeds probably did not limit rodent populations or distribution. If seed of these species were not the limiting factor, then ro- dent populations should not have declined by 58 percent in areas of high ant-density. One might suspect an error in our data due to in- appropriate sampling, but comparable results were obtained on the base grid. In August 1977 and 1978, significant shifts occurred in the centers of activity from the area of high ant-density to the area of low ant-density (Table 4). In fact, two P. manicitkitus that es- tablished centers of activity in the area of high ant-density during May established new centers of activity in the area of low ant-den- sity during August. The negative correlation between ants and 3.0 - • • (y= 4,329) 2.7 - ^ 2.4 - • • 2.1 - • 1.8 1.5 - • • 1.2 - • • -^ 0.9 - • • • • • •• 0.6 - • • •• 0.3 - 1.0 1.5 2.0 2.5 3.0 3.5 4.0 4.5 5.0 5.5 6.0 6.5 7.0 Estimated Number of Active Ants ix xlOO) Fig. 1. Regression analysis of mound volume and estimated number of active ants (p< 0.050). 270 Great Basin Naturalist Vol. 39, No. 3 Table 1. Relative seed abundance and time of seed drop for Descuniinia richardsonii, Lepidium perfoliutum, and Sitanion hystrix for May-August 1977. Annual seeds Grass seeds per sample per sample High ant- Low ant- High ant- Low ant- Date density density Average density density Average May 128 25 77 0 27 14 June 242 53 147 4 31 18 July 199 126 163 2 86 44 August 144 16 81 0 78 39 Average 179 55 3 56 Table 2. Numbers and relative percentage of seeds from Descuniinia ricliardsonii, Lepidium perfoliatuin and Sita- nion hystrix (grass) collected from 75 foraging ants at five ant mounds for the four sampling periods in 1978. Annual seeds Grass seeds Number combined Percent of total Date Annual seeds Grass seeds June 23 July 6 July 16 July 28 358 249 47 39 1 16 88 111 359 265 135 150 99.7 94.0 35.0 26.0 0.3 6.0 65.0 74.0 20 U) ^ 12 1000 3000 2000 Ant Density Fig. 2. The correlation between ant and rodent density (p< 0.025) is shown for May-August 1978. 4000 Sept. 1979 Landeen et al.: Ant-Rodent Competition 271 rodents (Fig. 2), the 58 percent rodent popu- lation decline in areas of high ant-density (Table 3), and the shifts in rodent centers of activity (Table 4) indicate that interactions did occur between ants and rodents. Com- petition for seeds is probably not responsible for these interactions; consequently, other factors such as rodent energetics, predation, and seed availability may logically be ex- pected to account for rodent distribution changes in areas of high ant-density. During the critical reproductive period in May, fe- male rodents must secure sufficient energy for existence and successful rearing of young. At that time it seems critical that females be in areas of relatively high food abundance in order to maximize reproductive energy. As- suming this to be the case, two predictions can be generated: (1) rodent numbers will be higher in areas where seed abundance is greatest and (2) sex ratios will be weighted toward females in areas of high seed-density. Both of these predictions were realized in our studies. Areas of high seed-density were also the areas of highest ant densities (Table 1), but, because ants were inactive in May (Table 3), competition between ants and ro- Table 3. Mean numbers ± standard deviation or ro- dents/grid in areas of high and low ant-densities, and the relative ant-foraging activity for the period May-August 1978. Mean no. rodents/grid Date High ant- density area Low ant- density area Relative foraging activity May June July August 26 ±8.3 29 ±6.2 17±4.1 11±2.7 9 ±3.2 24 ±4.3 18±2.8 16±2.7 8±12 56 ±30 104 ±70 26 ±13 dents for seeds was nonexistent at that time. Because of the temporal difference in seed drop between the annual herbaceous species (D. richardsonii, L. perfoliatwn) and gra.ss (S. hystrix), seeds from annual plants are more available in May and June, whereas the grass seeds are more available in July and August. This difference in seed availability was also reflected in the foraging activities of harves- ter ants. They foraged almost entirely on seeds from annual plants in June, but, as the season progressed and these seeds became less available due to ant-rodent foraging ac- tivities and the loss of seeds into soil crevices (Tevis 1958, Bernstein 1974), ants foraged primarily on grass seeds that were more available (Table 2). Since grasses were vir- tually nonexistent in areas of high ant-density (Table 1), rodents likely emmigrated to areas where those seeds were more abundant. Rodents living in areas of high ant-density would probably be more susceptible to pre- dators than those living in areas of low ant- density for two reasons: (1) areas of high ant- density have more open spaces and less total cover than areas of low ant-density (Table 1), (2) foraging time will increase as the seed availability of D. richardsonii and L. per- foliatum decrease. Observations in July and August indicated that predator pressure on rodents was greater in areas of high ant-den- sity. Short-eared owls and burrowing owls were observed almost exclusively in those areas. Coyote signs were more abundant in areas of high ant-density, and the only weasel captured during the summer was on a grid with high ant-density. Because ecological theory suggests that small animals are more likely to be regulated by predation than com- petition (Hairston et al. 1960, Schoener 1974), our results suggest an alternative ex- Table 4. Distribution of centers of rodent activity for areas of high and low ant-densities on the base grid for May and August 1977 and 1978. Rodent May August centers of activity High ant- density area Low ant- density area Total High ant- density area Low ant- density area Total Number, 1977 Number, 1978 Percentage, 1977 Percentage, 1978 16 31 37 48 27 34 63 52 43 65 3 16 20 35 12 29 80 65 15 45 272 Great Basin Naturalist Vol. 39, No. 3 ifl planation for observed ant-rodent distribu- tion patterns to that proposed by Brown and Davidson (1977). Differential predation pressure and seed availability are factors that have not been treated as complimentary mechanisms in- fluencing rodent distributions in areas of high ant-density. Although this is the first reported study of ant-rodent interactions in a cold desert, the results suggest that competition for seeds between the two taxa may not be as intense in cold deserts as indicated by Brown et al. (1975), Brown and Davidson (1977), and Reichman (unpubl. ms.) for the Sonoran Desert. Pulliam and Brand (1975) reported that competition for seeds between ants and rodents in the plains grassland region of Ari- zona may occur only in periods of drought. Because 1978 was an exceptional year for production of annuals (19.2 percent relative cover compared to 6.8 percent in 1977), ac- tive competition for seeds between the two taxa may have been masked. Ants may exert a stronger influence on ro- dent distributions in cold desert environments than our data have shown. Because ants have an impact on the total vegetation cover of an area (Rogers and Lavigne 1974, Clark and Comanor 1975), their absence may allow in- creases in total shrub and grass cover, thus af- fecting rodent distributions. The decrease in availability of seed from annual plants prob- ably would not be as dramatic if there were no ants present. If this were the case, then fo- raging time and consequent predation would not increase significantly as the season pro- gressed. Further documentation of selection pressures such as predation and seed avail- ability during drought years are needed be- fore the extent of competition between ants and rodents in cold desert environments can be fully evaluated. Literature Cited Bernstein, R. A. 1974. Seasonal food abundance and fo- raging activity in some desert ants. Amer. Nat. 108:490-498. BoYDEN, T. C. 1978. Territorial defense against hum- mingbirds and insects by tropical hummingbirds. Condor 80:216-221. Brown, J. H. 1971. Mechanisms of competitive exclu- sion between two species of chipmunks. Ecology 52:305-311. Brown, J. H., J. J. Grover, D. W. Davidson, and G. A. LiEBERMAN. 1975. A preliminary study of seed predation in desert and montane habitats. Ecolo- gy 56:987-992. Brown, J. H., and D. W. Davidson. 1977. Competition between seed-eating rodents and ants in desert ecosystems. Science 196:880-882. Clark, W. H., and P. L. Comanor. 1975. Removal of annual plants from the desert ecosystem by west- em harvester ants, Pogonomijnnex occidentalis. Environ. Ent. 4:52-56. Cole, A. C. 1932. The relationship of the ant, Pogono- mijnnex occidentalis Cresson, to its habitat. Ohio J. Sci. .32:133-146. . 19.34. An ecological study of the ants of the southern desert shrub region of the United States. Ann. Entomol. Soc. Amer. 27:388-405. . 1968. Pogonomijnnex harvest ants: a study of the genus in North America. Univ. Tenn. Press, Knoxville. 222 p. CoNNELL, J. H. 1961. The influence of interspecific com- petition and other factors on the distribution of the barnacle Chthamalus stellatiis. Ecology 42:710-723. Creighton, W. S. 19.53. New data on the habits of the ants of the genus Veromessor. Amer. Mus. Novit. 1612:1-18. Davidson, D. W. 1977a. Species diversity and commu- nity organization in desert seed-eating ants. Ecol- ogy 53:711-724. . 1977b. Foraging ecology and community organi- zation in desert seed-eating ants. Ecology 58:725-737. Fenton, M. B., and T. H. Fleming. 1975. Ecological in- teractions between bats and nocturnal birds. Biotropica 8:104-110. Franz, C. E., O. J. Reichman, and K. M. Van De Graff. 1973. Diets, food preferences, and repro- ductive cycles of some desert rodents. US/lBP/Desert Biome Research Memorandum RM 73-24 128 p. Garcia, J. R., H. D. Smith, and C. D. Jorgensen. 1974. A capture-release method for determining small mammal activity. Proc. Utah Acad, of Sci. Arts and Letters. 51:1-11. Grant, P. R. 1972. Interspecific competition among ro- dents. Ann. Rev. Ecol. & Syst. 3:79-106. Hairston, G. H., F. E. Smith, and L. B. Slobodkin. 1960. Community structure, population control, and competition. Amer. Nat. 94:421-425. Hansen, R. M., and D. M. Ueckert. 1970. Dietary sim- ilarity of some primary consumers. Ecology 51:640-648. Hayne, D. W. 1949a. Calculation of size of home range. J. Mamm. 30:1-18. . 1949b. Two methods for estimating population from trapping records. J. Mamm. 30:.399-411. Hull, A. C, Jr., and J. R. Killough. 1951. Ants are consuming Big Horn Basin ranges. Annual Live- stock Review, Herdsman Edition, Sec. 2, Western Farm Life 53:70. Lavigne, R. J. 1966. Individual mound treatments for control of the western harvester ant, Pogonomyr- Sept. 1979 Landeen et al.: Ant-Rodent Competition 273 niex occidciitalis, in Wyoming. J. Econ. Entoni. 59:525-5.32. Mares, M. A., and M. L. Rosenzweig. 1978. Cranivory in North and South American de.sert.s: rodents, birds and ants. Ecology 59:235-241. MACi\RTHUR, R. H. 1972. Geographical ecology. Harper & Row, New York. p. 111-207. PiANKA, E. R. 1974. Evolutionary ecology. Harper & Row, New York. p. 1.32-155. Prim.\ck, R. B., AND H. F. Howe. 1975. Interference competition between a hummingbird and skipper butterflies. Biotropica 7:55-58. [\jLLiAM, R. H., AND M. R. Brand. 1975. The production and utilization of seeds in plains grassland of southeastern Arizona. Ecology 56:1158-1166. Race, S. R. 1966. Control of western harvester ants on rangeland. N.M. Agric. Exp. Stn. Bull. 502. 21p. Reichman, O. J. 1979. Desert granivora foraging and its impact on seed densities and distributions (un- publ. ms). Rogers, L. E. 1974. Foraging activity of the western harvester ant in the shortgrass plains ecosystem. Environ. Entom. 3:420-424. Rogers, L. E., and R. J. Lavigne. 1974. Environmental effects of western harvester ants (Pogonomyrmex occidentalis) on the short grass plains ecosystem. Environ. Entom. 3:994-997. ScHOENER, R. W. 1974. Resource partitioning in ecologi- cal commimities. Science 185:27-.39. Schroder, G. D., and M. S. Rosenzweig. 1975. Per- turbation analysis of competition and overlap in habitat utilization between Dipodomijs ordii and Dipodotnys merriami. Oecologia 19:9-28. .Severin, H. C. 1955. Harvester ants and their control. S.D. Farm and Home Res. Winter 6:36-37. Sharp, L. A., and W. F. Barr. 1960. Preliminary in- vestigations of harvester ants on southern Idaho rangelands. J. Range Mgmt. 13:131-1.34. Sheppard, D. H. 1971. Competition between two chip- munk species (Eiitamias). Ecology 52:320-.329. Sinclair, A. R. E. 1975. The resource limitation of tro- phic levels in tropical grassland ecosystems. J. Anim. Ecol. 44:497-520. Tevis, L., Jr. 1958. Interrelations between the harvester ant, Veromessor pergandei (Mayr), and some desert ephemerals. Ecology .39:695-704. Whitford, W. G. 1976. Foraging behavior in Chihua- huan Desert harvest ants. Amer. Midi. Natur. 95:455-458. Whitford, W. G., and G. Ettershank. 1975. Factors affecting foraging activity in Chihuahuan Desert harvester ants. Environ. Entom. 4:689-696. Wiens, J. A. 1977. On competition and variable environ- ments. Amer. Sci. 65:590-597. Wight, J. R., and J. T. Nichols. 1966. Effects of harves- ter ants on production of a saltbush community. J. Range Mgmt. 19:68-71. FOOD HABITS OF BURROWING OWLS IN SOUTHEASTERN IDAHO R. L. Gleason' and T. H. Craig- Abstract.— Food habits of a population of the Burrowing Owl {Athene cunicularia) at the Idaho National Engi- neering Laboratory, Butte County, were studied. The 421 pellets examined yielded 2,436 prey items of at least 22 prey species. Invertebrates, largely insects, constituted 91 percent of the total prey items, but only 29 percent of the total biomass; mammals constituted 8 percent of the prey items, but 68 percent of the biomass. The prey were most- ly nocturnal species; diurnal species were poorly represented. The Burrowing Owl (Athene cunicularia) is widely distributed in the dry prairies and grasslands of western North America. In Idaho, Burrowing Owls are breeding resi- dents on the Snake River Plains, generally ar- riving in April and departing in October. Al- though the food habits of Burrowing Owls have been documented in much of their range (Robertson 1929, Errington and Ben- nett 1935, Hamilton 1941, Sperry 1941, Bond 1942, Longhurst 1942, Thomsen 1971, Butts 1973, Marti 1974), including Oregon (Maser et al. 1971) and Utah (Smith and Murphy 1973a), this paper provides the first informa- tion from Idaho. The study population was located within and adjacent to the 2315 km- Idaho National Engineering Laboratory (INEL) Site in Butte Co. Elevation on the INEL Site ranges from 1454 m to 1554 m. Rainfall averages less than 25 cm a year, with large annual variations. The average monthly temperatures are below 0 C for November through March, and ap- proach 20 C for the hottest month, July. Veg- etation is dominated by big sagebrush [Arte- misia tridentata), which covers about 80 percent of the site (Harniss and West 1973). Pellets were collected at nine active bur- rows from 4 June to 29 July 1975. This was approximately the period from hatching of the young to dispersal of the family groups. Burrows were visited about every four days and pellets were usually found within a radi- us of several meters from the burrow en- trances. Pellets were dissected in the labora- tory and the number and identity of prey remains in each pellet were recorded. Mam- malian remains were identified by dental characteristics, and mandibles were paired to determine the total number of individuals represented. Invertebrate prey were identi- fied and counted using recovered mandibles, heads, and/ or elytra. Average body weights were taken from Smith and Murphy (1973b), Evans and Emlen (1974), Johnson (1977), and Diller (pers. comm.). A total of 421 pellets yielded 2,436 prey items and indicated that the owls utilized at least 22 species of prey (Table 1). In- vertebrates were the most frequent prey of Burrowing Owls on the INEL Site, con- stituting about 91 percent of the total num- ber of prey items. However, invertebrates represented only 29 percent of the total bio- mass. Conversely, mammals represented 8 percent of the total prey items, and 68 per- cent of the total biomass. Similarly, Smith and Murphy (1973b) found that over two breeding seasons invertebrates constituted 80 percent of the total prey by number, but only 5 percent of the biomass. Marti (1974) found that over three seasons in Colorado in- vertebrates made up 90 percent of the prey by number, but only 9 percent of the bio- mass. Earhart and Johnson (1970) and Murray (1976), in theoretical discussions of sexual di- morphism and geographical variation in clutch sizes in North American owls, suggest 'Department of Zoology, University of Idaho, Moscow, Idaho 83843. 'Route 3, Box 170, Cottonwood, Idaho 83255. 274 Sept. 1979 Gleason, Craig: Burrowing Owls 275 that Burrowing Owls are primarily in- sectivorous. Though this conclusion can be drawn from reports on food habits which rank the relative importance of prey only in terms of percent of the total of prey items, it does not appear justified when considering the prey composition in terms of total bio- mass. For predators such as Burrowing Owls which utilize prey differing greatly in size, biomass estimates should be included to pre- sent an accurate picture of diet composition. Sixty-six percent of the total prey biomass of Burrowing Owls on the INEL Site was represented by three species, Ord kangaroo rat {Dipodmnys ordii), montane vole {Micro- tus montaniis), and Jenisalem cricket {Steno- pehnatus ftiscus). All of the burrows we stud- ied were probably originally excavated by badgers {Taxidea taxus) in pursuit of either kangaroo rats or Great Basin pocket mice (Perognathus parvus). Thus all the burrows were in or near established colonies of these species. In addition, seven of the burrows were in areas adjacent to cultivated fields of alfalfa {Medicago sativa), which frequently harbor large populations of montane voles. Jerusalem crickets, which composed almost half of the prey by number, are extremely common insects in arid regions of western North America. They are large, slow, and conspicuous, traits which make them very vulnerable to predation by Burrowing Owls. From this study it would appear that Bur- rowing Owls in southeastern Idaho are pri- marily nocturnal predators. All of the mam- mal and the majority of invertebrate prey of the owls are most active at night. The most frequently occurring prey species, Jerusalem crickets, are strictly nocturnal. Grasshoppers and passerine birds, both principally diurnal, contributed little to the total diet, even though both were abundant in the area. Table 1. Food habits of Burrowing Owls in southeastern Idaho as determined by pellet analysis. Number Percent Biomass Percent (g) biomass (6882) (68) 2420 24 2052 20 900 9 595 6 900 9 15 tr (140) (1) 84 1 56 tr Mammals Dipodomys ordii Microtus montanus Perognathus parvus Peromyscus maniculatus Thomomys talpoides Mus musculus Birds Eremophila alpestris Unidentified passerine Amphibians Scaphiopus interrnontanus Arachnids Scorpionidae Solpugidae Insects Gryllacrididae Acrididae Cicadidae Carabidae Silphidae Scarabaeidae Tenebrionidae Formicidae Unidentified Coleoptera Totals (200) 44 54 60 35 (10) 6 4 (8) 2 2 3 1 tr tr (tr) tr tr (283) (12) (338) (4) 123 5 178 2 160 7 160 2 (1924) (79) (2501) (25) 1122 46 2244 22 8 tr 5 tr 29 1 29 tr 62 3 14 tr 133 5 40 tr 129 5 39 tr 42 2 23 tr 61 3 6 tr 338 14 101 1 2436 10.127 276 Great Basin Naturalist Vol. 39, No. 3 We thank D. R. Johnson, University of Idaho, for reviewing the manuscript, and W. F. Barr, University of Idaho, for his very gen- erous contribution of time and expertise in identifying innumerable insect fragments. This research was funded by the Division of Biomedics and Environmental Research (DOE) and is a contribution of the INEL Site Ecology Studies Program. Literature Cited Bond, R. M. 1942. Food of the Burrowing Owl in west- ern Nevada. Condor 44:183. Butts, K. O. 1973. Life history and habitat requirements of Burrowing Owls in western Oklahoma. Unpub- lished thesis, Oklahoma State University, Stillwa- ter, Oklahoma. 188 p. Earhart, C. M., and N. K. Johnson. 1970. Size di- morphism and food habits of North American owls. Condor 72:251-264. Errington, p. L., and J. L. Bennett. 1935. Food habits of Burrowing Owls in northwestern Iowa. Wilson Bull. 47:125-128. Evans, F. C, and J. T. Emlen, Jr. 1947. Ecological notes on the prey selected by a Barn Owl. Con- dor 49:.3-9. Hamilton, W. J. 1941. A note on the food of the west- ern Burrowing Owl. Condor 43:74. Harniss, R. O., and N. E. West. 1973. Vegetation pat- terns of the National Reactor Testing Station, southeastern Idaho. Northwest Sci. 47;.30-43. Johnson, W. C. 1977. Examination of consuming tech- niques for small mammals in a high desert ecosys- tem. Task No. 8, Idaho National Engineering Site Ecological Studies, Idaho Falls, Idaho. 95 p. LoNGHURST, W. M. 1942. The summer food of Bur- rowing Owls in Costilla Coimtv, Colorado. Con- dor 44:281-282. Marti, C. D. 1974. Feeding ecology of four sympatric owls. Condor 76:45-61. Maser, C. E., E. W. Ham.mer, and S. H. Anderson. 1971. Food habits of the Burrowing Owl in cen- tral Oregon. Northwest Sci. 45:19-26. Murray, G. A. 1976. Geographic variation in the clutch .sizes of several owl species. Auk 93:602-613. Robertson, J. 1929. Some observations on the feeding, habits of the Burrowing Owl. Condor 31:.38-.39. Smith, D. G., and J. R. Murphy. 1973a. Late summer food habits of adult Burrowing Owls in central Utah. Raptor Research 7:112-115. . 1973b. Breeding ecology of raptors in the eastern great Basin of Utah. Brigham Young Univ. Sci. Bull., Biol. Ser. 18(3): 1-76. Sperry, C. C. 1941. Burrowing Owls eat spadefoot toads. Wilson Bull. 53:45. Thomsen, L. 1971. Behavior and ecology of Burrowing Owls on the Oakland Municipal Airport. Condor 73:177-192. FLORA OF THE LEE CREEK VALLEY, ALBERTA R. Keith Shaw' ,\bstract.— The floristic composition of the Lee Creek valley in southwestern Alberta, Canada, is presented. The valley flora consists of 299 species of vascular plants representing 173 genera and 46 families. The flora is dominated by forbs (73 percent), followed by grasses (24 percent), shrubs (12 percent), and trees (11 percent). The valley of Lee Creek is divided into upper and lower sections on the basis of elevational, climatic, and biotic differences. Floristic richness of the Lee Creek valley is quite evenly divided between the upper and lower valleys: 105 species limited to the upper valley, 95 to the lower valley, and 99 species .shared by both upper and lower valleys. In this paper the results of my field and herbarium work over the years 1957-1978 are presented for that part of the Lee Creek valley lying inside the southwestern corner of the Province of Alberta. The ecological as- pects of the river bottom forest biome along the prairie section of Lee Creek have been reported (Shaw 1976), with a combined plant species list for the river bottom forest on St. Mary River, Lee Creek, and Belly River. This paper borrows the river bottom forest species list for lower Lee Creek and adds to it the greater species lists for the other Lee Creek valley biomes. Geology and Geography Lee Creek originates on the north and east slopes of Old Chief Mountain in Glacier Na- tional Park, Montana. It flows from alpine meadows at 2,000-3,000 m in three branches that join where the creek crosses the Inter- national Boundary at Alberta's Range 27. The creek then flows through the lower montane forest and aspen parkland biomes of the up- per valley to Section 11 Township 2 Range 27 West of 4, where it crosses the treeless stretches of the fescue prairie biome at eleva- tions of 1,400 m down to 900 m. Along the lower stream course the poplar-dominated river bottom forest biome becomes a unique ecological entity on Lee Creek. This biome's presence identifies the lower valley and is continuous downstream, through the town of Cardston, to the creek's confluence with St. Mary River at Section 23 Town.ship 3 Range 25 West of 4. Lee Creek drains about 290 square kilometers of northwest Montana and southwest Alberta. Streamside and valley floor vegetation pat- terns are modified by Lee Creek streamflow (Shaw 1976). Monthly water flow varies widely throughout the year (Table 1). From late July through autumn and winter the flow is fairly constant and moderate, but during March warmer weather causes snow melt in the foothills and on the lower mountain slopes to increase stream flow. The most rap- id melting of deep mountain snow occurs in late May and early June. This coincides with the season of highest precipitation (Table 2), swelling stream flow to its maximum, which is four to five times the winter flow rate. Se- vere flooding, with considerable streamside vegetation and habitat alteration, occurred in 1902, 1908, 1950, 1964, and 1975. The mountain section of the Lee Creek valley, in Montana, is carved through geolog- ical formations of the Lewis Range's Belt Series and transported eastward as the Lewis Overthrust and overlying the younger Cre- taceous shales and sandstones of the plains (Wyatt 1939). The prairie section of Lee Creek flows through and over a variety of consolidated and unconsolidated deposits, from the transported Belt series rocks of Pro- terozoic time to the more recent Cretaceous and Tertiary series. Bedrock cut through and 'Box 364, Cardston, Alberta TOK OKO. 277 278 Great Basin Naturalist Vol. 39, No. 3 exposed along the Alberta section of Lee Creek is mainly an exposure of the highly calcareous light gray sandstones and sandy shales of the St. Mary River nonmarine sand- stone, the uppermost of the Cretaceous for- mations of southwest Alberta. Irregular bed- ding and cross-bedding are common, and freshwater oyster shells and coal beds are fre- quently exposed. Much of this bedrock is ve- neered with glacial deposits left when the Ice Age glaciers melted. There is also widespread distribution of reworked glacial deposits as well as alluvial and lacustrine deposits trans- ported by the creek. The soils along the creek valley bottom are of alluvial deposition and some, still liable to frequent flooding, are quite variable in ferti- lity, texture, and utilization. In southwest Alberta the break from mountains to plains is fairly rapid, there being no wide range of foothills. Climate Summer along Lee Creek is normally warm, but winter is usually long and cold (Table 3). Mean temperatures are below 0 degrees C from November through March at Cardston on lower Lee Creek. A wide varia- tion in winter temperatures occurs, depen- ding on the southerly flow of cold Arctic air and the easterly flow of temperature-moder- ating Chinook winds. Temperatures rise rap- idly from winter to summer and decline with equal rapidity from summer to winter. Monthly mean temperatures at Cardston are above 10 degrees C for the five months from May to September. The average frost-free period on Lee Creek to which the native plants have adapted ranges from 70 days in montane forest on upper Lee Creek to 100 days at Cardston near the creek's confluence. Precipitation along the valley shows wide variation from year to year. Winter snowfall in the lower valley is comparatively light, with amounts ranging from 76 to 127 cm. This increases in the upper valley to 180 cm. Total precipitation at Cardston in the lower valley averages 45.8 cm and gradually in- creases through the upper valley (Canada Transport 1967). A combination of snowfall and wind causes drift buildup in the stream valley throughout winter. Ecology and Taxonomy The Lee Creek valley in Alberta is a post- glacial, meandering, stream-carved valley. The stream itself works back and forth from Table 1. Mean monthly stream flow in cubic meters per second of Lee Creek in southwest Alberta, Canada. Month Lee Creek Cardston January February March April May June July August September October November December 0.4 0.4 1.1 2.4 4.6 4.5 1.5 0.5 0.5 0.6 0.5 0.3 Table 2. Mean monthly precipitation in inches and centimeters for Cardston, Alberta, elevation 1,151 m. Month Inches Centimeters January February 0.75 0.91 1.9 2.3 March 1.14 2.9 April May June 1.39 2.38 3.59 3.5 6.1 9.1 July August September October 1.62 1.50 1.75 1.19 4.1 3.8 4.4 3.0 November 0.94 2.4 December 0.88 2.2 Total 18.04 45.8 Table 3. Mean daily temperature in degrees Centi- grade and degrees Fahrenheit at Cardston, Alberta. Month Centigrade Fahrenheit January -7.4 18.7 February -6.5 20.3 March -2.5 27.5 April 4.5 40.2 May 10.3 50.6 June 13.7 56.5 July 17.9 64.1 August 16.1 61.1 September 11.6 52.7 October 6.6 43.8 November -0.4 31.3 December -4.2 24.4 Year 5.0 40.9 Sept. 1979 Shaw: Alberta Flora 279 year to year resorting its gravel base, eroding on the outside bends and depositing on the inside ones as it cuts a deeper channel. Major alteration occms during the season of high water in late May and early June. The life span of a gravel bar, the first ter- race of the valley floor, is entirely fortuitous, depending on the vagaries of Alberta weath- er and its effect on streamflow. If the first terrace escapes erosional destruction for 20 years or more, natural succession will change the herb-dominated first terrace plant com- mimity to the poplar-dominated lower val- ley, and poplar and spruce-dominated upper valley, communities on the terrace. As ero- sion and channel shifting are directed away from the terrace and the stream cuts deeper into the valley floor, what was once a first terrace becomes a second terrace, and it is here the river bottom forest thrives. Years or centuries later the second terrace may be left even higher as a third terrace, and when this happens the river bottom forest fails to be self-sustaining. As the mature trees die out without seed or vegetative replacement, the grassland biome (fescue prairie) in the lower valley, or in the upper valley the deciduous forest biome (aspen parkland), take over with virtual similarity to the vegetation on valley slopes and surrounding terrain. With the constant erosion, deposition, and channel shifting, fresh bars of material sorted into various textures and of varying depths annually become available for colonization by plants. In spite of this constancy of change, the creek bottom floodplain is a very uniform habitat where climate and substrate determine the establishment of plant species. The combined upper and lower Lee Creek vascular flora consists of 49 species of woody plants and 250 species of herbaceous ones, for a total of 299 species. In ecological distribution 105 species were confined to the upper Lee Creek valley, 95 species to the lower Lee Creek valley, and 99 species were found in both the upper and lower valleys. Naturalized exotics made up 7.7 percent (23 species) and native species 92.3 percent (276 species) of the total species number. Taxonomically the 299 species were con- -tributed by 46 families and 173 genera. Nine families dominated the flora. A dominant family arbitrarily was one with 10 or more species in the valley (Table 4). Floristic rich- ness is enhanced by the high yearly frequen- cy of new gravel bar pioneer sites and the availability of plant reproductive material from five major biomes: alpine tundra, mon- tane forest, aspen parkland, fescue prairie, and river bottom forest. Within the Lee Creek valley flora there are 11 plant species listed by Argus and White (1978) as being rare in Alberta. These 11 are Angelica dawsonii, Balsamorhiza sa- gittata, Calochortus apiciilatus, Disporum oreganum, Hydrophyllum capitatwn, Larix occidentalis, Lesquerella alpina, Phlox alyssi- folia, Plantago canescens, Populus angusti- folia, and Senecio hydrophiloides. Lee Creek, like many other stream sys- tems, provides excellent seed dispersal habi- tat. Flowing water can bring mountain or submontane species to greater range exten- sions down the valley. Windborne seeds are readily dropped in the lee of sheltering banks and thickets as wind velocity decreases. Bird activities provide other transport mecha- nisms. Downstream species are able to extend their ranges upstream nearly as readily (Table 5). All parts of the Lee Creek valley in Al- berta are grazed by domestic livestock, prin- cipally cattle. The upper valley represents mainly summer-use pasture and the lower valley receives year-round grazing use. Deer, both white-tail and mule, are common throughout the length of the valley. Intensive plant collecting was done at many valley sites throughout the growing Table 4. Dominant plant families of the Lee Creek valley. Number Percent of Family of species total species Compositae 52 17.4 Leguminosae 32 10.7 Gramineae 24 8.0 Liliaceae 17 5.7 Rosaceae 17 5.7 Salicaceae 15 5.0 Ranunculaceae 13 4.3 Scrophulariaceae 12 4.0 Orchidaceae 10 3.3 Total 192 64.2 280 Great Basin Naturalist Vol. 39, No. 3 seasons of several years. The valley flowering season usually began in early May and ended by late September. Specimens were pre- served according to standard herbarium prac- tice. Almost all identifications were checked against known herbarium reference material. Taxonomy of the poplars follows Brayshaw (1965), the genus Cryptantha after Higgins (1971), and the remainder after Moss (1959), Boivin (1969), Booth (1950), and Booth and Wright (1966). The willows were identified by George Argus of the National Museum of Natural Sciences. Only those flowering plants native to Lee Creek valley, or naturalized and viable exot- ics, are included in the species list. Cultivated agricultural and ornamental species were de- leted. The valley is considered to be all the area below the rim elevation of the adjoining land surface. All specimens collected are in the author's personal herbarium at Cardston, Alberta. A nearly complete duplicate set is on deposit in the Brigham Young University Herbarium (BRY) at Provo, Utah. For convenience, plant families, genera, and species are listed in alphabetical order. Because of the ecological differences along Lee Creek between the upper and lower val- leys, plant species listed will be designated as found in the upper valley only (U), lower val- ley only (L), or in both upper and lower val- leys (UL). Species List Berberidaceae Berberis repens Lindl. U Betulaceae Betula glandulosa Michx. U Betula occidentalis Hook. UL Betula pumila L. var. glandulifera Kegel U BORAGINACEAE Hackelia floribunda (Lehm.) I.M. Johnston UL Lappula echinata Gilib. L Lithospermtim incisuin Lehni. L Lithospennum rudende Lehm. UL Campanulaceae Campanula rotundifolia L. UL Capparidaceae Cleome serrtdata Pursh L Caprifoliaceae Linnaea borealis L. var. americana (Forbes) Rehd. U Lonicera dioica L. var. glaucescens (Rydb.) Butters UL Lonicera involucrata (Richards.) Banks U Lonicera tartarica L. L Sytnphoricarpos occidentalis Hook. UL Viburnum edule (Michx.) Raf. U Caryophyllaceae Arenaria lateriflora L. UL Arenaria rubella (Wahlenb.) J.E. Sin. U Cerastium arvense L. UL Silene menziesii Hook. U Stellaria longipes Goldie U COMPOSITAE Achillea millefolium L. UL Agoseris glauca (Pursh) Raf. UL Anaphalis margaritacea (L.) Benth. & Hook. U Antennaria neglecta Greene U Antennaria pulchcrrima (Hook.) Greene U Antennaria racemosa Hook. U Antennaria rosea Greene UL Arnica chamissonis Less. ssp. foliosa (Nutt.) Mag. U Arnica cordifolia Hook. U Table 5. Dispersal of species up and down the Lee Creek valley beyond their usual geographic ranges. Downstream range extensions Upstream range extensions Actaea rubra Arctostaphylos uva-ursi Astragalus bourgovii Corallorhiza striata Epilobium angustifolium Epilobiwn latifolium Heracleum lanatum Larix occidentalis Picea glauca Pinus flexilis Populus balsamifera trichocarpa Populus tremuloides Pseudotsuga menziesii Pyrola asarifolia purpurea Smilacina racemosa amplexicaulis Veratrum eschscholtzii Astragalus flexuosus Elaeagnus commutata Oxytropis campestris gracilis Oxytropis sericea spicata Oxytropis viscida Populus angustifolia Salix interior Sept. 1979 Shaw: Alberta Flora 281 Arnica lonchophylla Greene U Arnica sororia Greene UL Artemisia biennis Willd. L Artemisia campestris L. L Artemisia ludoviciana Nutt. L Aster foliaceiis Lindl. U Aster kievis L. var. geycri A. Gray UL Aster occidentalis (Nutt.) T. & G. UL Aster pansus (Blake) Cronq. UL Aster sibiricus L. U Balsamorhiza sagittata (Pursh) Nutt. UL Chrysanthemum leucanthemwn L. UL Chrysopsis villosa (Pursh) Nutt. var. hispida (Hook.) Gray L Cichorium intybiis L. L Cirsium arvense (L.) Scop. UL Cirsium undulatum (Nutt.) Spreng. L Crepis elegans Hook. U Erigeron caespitosus Nutt. L Erigeron glabellus Nutt. var. pitbescens (Hook.) Cronq. U Erigeron peregrinus (Pursh) Greene ssp. calianthemus (Greene) Cronq. L Erigeron philadelphicus L. U Erigeron speciosiis (Lindl.) DC. L Erigeron strigosus Muhl. L Erigeron stibtrinervis Rydb. var. conspicuus (Rydb.) Cronq. U Gaillardia aristata Pursh UL Helianthus laetiflorus Pers. var. subrhomboideus (Rydb.) Fern. L Hieracium scouleri A A. U Hieracium umbellatiim L. L Hyvienoxys acaulis (Pursh) Parker L Liatris punctata Hook. L Ratibida columnifera (Nutt.) Wooton & Standi. L Rudbeckia serotina Nutt. U Senecio canus Hook. UL Senecio hydrophiloides Rydb. U Senecio integerrimus Nutt. var. exaltatus (Nutt.) Cronq. L Senecio lugens Richards. UL Senecio pauperculus Michx. U Senecio triangularis Hook. U Solidago decumbens Greene var. oreophila (Rydb.) Fern. L Solidago missouriensis Nutt. UL Solidago miiltiradiata Ait. U Taraxacum officinale Weber UL Tragopogon dubius Scop. L CORNACEAE Cornus canadensis L. U Cornus stolonifera Michx. UL Crassulaceae Sedum stenopetalum Pursh U Cruciferae Arabis divaricata A. Nels. U Arabis hirsuta (L.) Scop. var. glabrata T. & G. UL Arabis holboellii Hornem. var. retrofracta (Graham) Rydb. L Erysimum cheiranthoides L. U Erysimum inconspicuum (S. Wats.) MacM. U Lesquerella alpina (Nutt.) S. Wats. var. spathulata - (Rydb.) Payson L Physaria didymocarpa (Hook.) A. Gray U Rorippa nasturtium-uquuticum (L.) Schinz & Thell. L Cyperaceae Carex filifolia Nutt. UL Carex hystricina Muhl. U Carex rostrata Stokes L Scirpus microcarpus Presl L Scirpus validus Vahl U Elaeagnaceae Elaeagnus commutata Bernh. UL Shepherdia argentea Nutt. L Shepherdia canadensis (L.) Nutt. U Equisetaceae Equisetum arvense L. UL Equisetum fluviatile L. U Ericaceae Arctostaphylos uva-ursi (L.) Spreng. UL Euphorbiaceae Euphorbia esula L. L Gentianaceae Gentiana detonsa Rottb. L Gentianella amarella (L.) Borner ssp. acuta (Michx.) J. M. Gillett U Geraniaceae Geranium nervosum Rydb. U Geranium richardsonii Fisch. & Trautv. U Geranium viscosissimum Fisch. & Mey. U Gramineae Agropyron smithii Rydb. L Agropyron subsecundum (Link.) Hitchc. L Agrostis alba L. L Beckmannia syzigachne (Steud.) Fern. L Bouteloua gracilis (HBK). Lag. L Brotnus commutatus Schrad. L Bromus inermis Leyss. L Calamagrostis purpurascens R. Br. U Danthonia parryi Scribn. U Danthonia spicata (L.) Beauv. U Deschampsia caespitosa (L.) Beauv. U Festuca idahoensis Elmer L Festuca ovina L. L Festuca scabrella Torr. UL Glyceria grandis S. Wats. UL Helictotrichon hookeri (Scribn.) Henr. U Koeleria cristata (L.) Pers. L Oryzopsis hymenoides (R. & S.) Ricker L Phleum pratense L. UL Poa arctica R. Br. U Poa compressa L. U Poa pratensis L. UL Stipa Columbiana Macoun U Stipa richardsonii Link U Hydrophyllaceae Hydrophyllum capitatum Dougl. U Phacelia sericea (Graham) A. Gray U Iridaceae Sisyrinchium montanum Greene U JUNCACEAE Juncus alpinus Vill. var. rariflorus Hartm. L Juncus longistylis Torr. L Labiatae Mentha arvensis L. var. villosa (Benth.) S.R. Stewart UL Monarda fistulosa L. var. menthaefolia (Graham) Fern. UL 282 Great Basin Naturalist Vol. 39, No. 3 Prunella vulgaris L. UL Leguminosae Astragalus aboriginum Richards. UL Astragalus alpinus L. U Astragalus bisulcatus (Hook.) A. Gray L Astragalus bourgovii A. Gray UL Astragalus canadensis L. L Astragalus crassicarpus Nutt. L Astragalus drummondii Dougl. L Astragalus flexuosus Dougl. UL Astragalus missouriensis Nutt. L Astragalus robinsii (Oakes) Gray UL Astragalus striatus Nutt. L Astragalus triphyllus Pursh L Astragalus vexilliflexus Sheld. U Glycyrrhiza lepidota (Nutt.) Pursh L Hedysarum alpinum L. UL Hedysarum sulphurescens Rydb. U Lathyrus ochroleucus Hook. UL Lupinus argenteus Pursh L Lupinus sericeus Pursh U Medicago lupulina L. L Medicago sativa L. L Melilotus alba Desr. L Melilotus officinalis (L.) Lam. L Oxytropis campestris (L.) DC. var. gracilis (A. Nels.) Barneby UL Oxytropis sericea Nutt. var. spicata (Hook.) Barneby UL Oxytropis splendens Dougl. UL Oxytropis viscida Nutt. UL Petalostemon purpureum (Vent.) Rydb. L Ttiemiopsis rhonibifolia (Nutt.) Richards. UL Trifolium hybridum L. L Trifolium pratense L. L Vicia americana Muhl. UL LiLIACEAE Allium cernuum Roth UL Allium schoenoprasum L. var. sibiricum (L.) Hartm. U Allium textile Nels. & Macbr. L Calochortus apiculatus Baker U Camassia quamash (Pursh) Greene U Clintonia uniflora (Schult.) Kunth U Disporum oreganum (S. Wats.) B. & H. L Erythronium grandiflorum Pursh U Fritillaria pudica (Pursh) Spreng. UL Lilium philadelphicum L. var. anditiurn (Nutt.) Ker U Smilacina racemosa (L.) Desf. var. amplexicaulis (Nutt.) S. Wats. UL Smilacina stellata (L.) Desf. UL Stenanthium occidentale A. Gray U Streptopus amplexifolius (L.) DC. U Veratrum eschscholtzii A. Gray U Zygadenus elegans Pursh UL Zygadenus gramineus Rydb. L LiNACEAE Linum lewisii Pursh UL Malvaceae Sphaeralcea coccinea (Pursh) Rydb. L Onacraceae Epilobium angustifolium L. U Epilobium glandulosum Lehm. L Epilobium latifolium L. UL Gaura coccinea Pursh var. glabra (Lehm.) Torr. & Gray L Oenothera biennis L. L Oenothera caespitosa Nutt. L Orchidaceae Calypso bulbosa (L.) Oakes U Corallorhiza maculata Raf. U Corallorhiza striata Lindl. UL Corallorhiza trifida Chatelain U Cypripedium passerinum Richards. U Habenaria dilatata (Pursh) Hook. U Habenaria obtusata (Pursh) Richards. U Habenaria unalascensis (Spreng.) S. Wats. U Habenaria viridis (L.) R. Br. var. bracteata (Muhl.) A. Gray UL Orchis rotundi folia Banks U Pinaceae Juniperus communis L. var. depressa Pursh UL Juniperus horizontalis Moench UL Larix occidentalis Nutt. U Picea glauca (Moench) Voss UL Pinus contorta Loudon var. latifolia Engelm. U Pinus flexilis James UL Pseudotsuga menziesii (Mirb.) Franco UL Plantaginaceae Plantago major L. UL Plantago septata Morris L = P. canescens Adams Polemoniaceae Collomia linearis Nutt. U Phlox alyssifolia Greene UL Phlox hoodii Richards. UL Polemonium pulcherrimum Hook. L POLYGONACEAE Eriogonum flavum Nutt. L Polygonum bistortoides Pursh U Rumex crispus L. L Primulaceae Dodecatheon conjugens Greene UL Dodecatheon radicatum Greene U Lysimachia ciliata L. UL Pyrolaceae Moneses uniflora (L.) A. Gray U Pyrola asarifolia Michx. var. purpurea (Bunge) Fern. UL Pyrola virens Schweigg. U Ranunculaceae Actaea rubra (Ait.) Willd. UL Anemone multifida Poir. UL Anemone parviflora Michx. U Anemone patens L. var. wolfgangiana (Bess.) Koch UL Aquilegia flavescens S. Wats. UL Clematis ligusticifolia Nutt. L Clematis verticellaris DC. var. columbiana (Nutt.) A. Gray UL Delphinium bicolor Nutt. UL Ranunculus acris L. UL Ranunculus flammula L. var. filiformis (Michx.) Hook. UL Ranunculus pedatifidus J.E. Smith var. affinis (R. Br.) L. Benson U Ranunculus sceleratus L. var. multifidus Nutt. L Tlialictrum venulosum Trel. U Rhamnaceae Rhamnus cathartica L. L Sept. 1979 Shaw: Alberta Flora 283 ROSACEAE Amelandiier alnifolia Nutt. UL Crataegus chnjfiocarf)a Ashe L Dryas dmminondii Richards. I'L Fragaria virgiiiimui Duchesne var. glaiica S. Wats. UL Geum allcpicuin Jaccj. UL Getim rivale L. U Geum triflorum Pursh UL Potentilla concinna Richards. L Potentilla fruticosa L. UL Potentilla glandulosa Lindl. ssp. pseudorupestris (Rydb.) Keck U Potentilla graeilis Dougl. var. flahelliformis (Lehni.) Nutt. U Potentilla gracilis Dougl. var. pulcherriina (Lehm.) Fern. U Prunus virginiana L. var. melanocarpa (A. Nels.) Sarg. UL Rosa acicularis Lindl. UL Rosa woodsii Lindl. UL Ruhus parviflorus Nutt. U Ruhus strigosus Michx. UL RUBIACEAE Galium boreale L. UL Galium triflorum Michx. L Salicaceae Popidus acuminata Rydb. L Populus angustifolia James L Popidus angustifolia James X halsamifera L. L Populus balsamifera L. subsp. halsamifera L Populus balsamifera L. subsp. trichocarpa (T.&G. ex Hook.) Bray.shaw UL Populus fremuloides Michx. UL Salix barclaiji Anderss. U Salix bebbiana Sarg. L Salix discolor Muhl. L Salix glauca Anderss. U Salix interior Rowlee L Salix lasiandra Benth. L Salix lutea Nutt. L Salix melanopsis Nutt. L Salix pseudomonticola Ball L Santalaceae Comandra pallida A. DC. UL Saxifragaceae Parnassia palustris L. var. neogaea Fern. L Ribes lacustre (Pars.) Poir. U Scrophulariaceae Besseija wijomingensis (A. Nels.) Rydb. L Castilleja miniata Dougl. UL Castilleja septentrionalis Lindl. U Orthocarpus luteus Nutt. L Pedicularis bracteosa Benth. U Pedicularis groenlandica Retz. U Penstemon confertus Dougl. UL Penstemon nitidus Dougl. UL Rhinanthus crista-galli L. U Verbascum thapsus L. L Verbena bracteata Lag. & Rodr. U Veronica americana (Raf.) Schw. UL Umbelliferae Angelica dawsonii S. Wats. U Ciciita douglasii (DC.) Coult. & Rose UL HeracU'um lanatum Michx. UL Lomatium simplex (Nutt.) Macbr. var. leptophyllum (Hook.) Mathias L Lomatium triternatum (Pursh) Coult. & Rose U Osmorhiza depauperata Phillippi U Perideridia gairdneri (Hook. & Am.) Mathias UL Zizia aptera (A. Gray) Fern. UL Valerianaceae Valeriana septentrionalis Rydb. U Violaceae Viola adunca J.E. Smith UL Viola nephrophylki Greene U Viola orbiculuta Geyer L Viola rugulosa Greene UL Viola selkirkii Pursh U Literature Cited Argus, G. and D. J. White. 1978. The rare vascular plants of Alberta. Syllogeus No. 17, National Mu- seum of Canada, Ottawa. BoiviN, B. 1969. Flora of the prairie provinces I, II, III, IV. Canada Dept. of Agriculture, Ottawa. Booth, W. E. 1950. Flora of Montana: I. Conifers and Monocots. Montana State Univ., Bozeman. Booth, W. E., and J. C. Wright. 1966. Flora of Mon- tana: II. Dicots. Montana State Univ., Bozeman. Brayshaw, T. C. 1965. Native poplars of southern Al- berta and their hybrids. Dept. Forestry Pub. No. 1109, The Queen's Printer, Ottawa. Canada Department of Transport. 1967. Temper- ature and precipitation tables for prairie provin- ces III. Ottawa. HiGGiNS, L. 1971. A revision of Cryptantha; subgenus Oreocarya. Brigham Young University Sci. Bull., Biol. Ser. 13 (4). Moss, E. H. 1959. Flora of Alberta. University of To- ronto, Ontario. Shaw, R. K. 1976. A taxonomic and ecologic study of the riverbottom forest on St. Mary River, Lee Creek and Belly River in southwestern Alberta, Canada. Great Basin Nat.: Vol. 36, No. 3. Wyatt, F. a. 1939. Soil survey of Lethbridge and Pinch- er Creek Sheets. University of Alberta, Edmon- ton. CLIMATES OF FESCUE GRASSLANDS OF MOUNTAINS IN THE WESTERN UNITED STATES T. Weaver' Abstract.— Climates of fescue grasslands were described by summarizing USDC Weather Bureau records of rep- resentative sites. Mean temperatures of the warmest month declined from 18 to 14 C, average annual precipitation increased from 40-50 to 170 cm, and the number of arid months in an arbitrarily defined dry year declined from 3 to 1-2 as one moves from Festuca arizonica (14 stations) to F. scabrella (15) to F. idahoensis (17) to F. thurberi (6) to F. viridula (9). Climates of the grasslands are sufficiently like those of the Pseudotsuga menziesii and Abies lasiocarpa zones of the northern Rocky Mountains to make one ask what factors— wind, snow duration, soil characteristics, or fire— allow the fescue grasslands to persist in a conifer climatic zone. Fescue grasslands form an important part of mountain vegetation in the western United States. The geographical and altitudinal ranges of major species are shown by Hitch- cock (1950) and Kuchler (1964) among oth- ers. Festuca arizonica Vasey occurs in the southern Rocky Moimtains under pine forests or in meadows in Kuchler 's types 18 and 19. Festuca thurberi Vasey is found in higher for- ests to timberline in the southern and central Rocky Mountains. Festuca scabrella Torr. and Festuca idahoensis Elmer occur in the northern Rocky Mountains in Kuchler's foot- hill grassland (63) type as well as in meadows in the forest zone. Festuca viridula Vasey is generally found west of the Rocky Mountains in alpine and subalpine meadows, above the ranges of Festuca scabrella and Festuca ida- hoensis. The objects of this paper are (1) to summa- rize data available for USDC Weather Bu- reau stations (6 to 17) in each grassland type and (2) to compare the climates of the fescue grasslands. Climates of adjacent vegetation types have been summarized by similar methods (Weaver 1979). Other studies of the climates of fescue grasslands include a 3-year study of summer climates of two Festuca sca- brella grasslands in British Columbia (van- Ryswyk et al. 1966), a 5 (now 10-)-year study of growing season climates of four Festuca idahoensis grasslands in southwest Montana (Mueggler 1971), and a 23-year study of cli- mate effects in a Festuca idahoensis stand in south Idaho (Blaisdell 1958). Methods Maps of weather station locations were sent to U.S. Forest Service and University personnel familiar with fescue grasslands, along with letters asking each to identify weather stations which lay in that person's fescue type. I gratefully acknowledge the help of E. Aldon, W. Clary, and W. Moir with F. arizonica stations; of P. Currie, W. Moir, H. Paulsen, and G. Turner with F. thurberi stations; of M. Morris with F. sca- brella stations; of M. Morris, W. Mueggler, and G. Payne with F. idahoensis stations; and J. Strickler with F. viridula stations. A few of the sites studied were not visited by those recommending them, but the availability of herbarium material from the sites and the small between-site standard errors (less than the between-year standard errors for a station in the type) suggest that there were few or no misclassifications. The 1961-1970 climatological data (USDC 1961-1970) for each station were summa- rized by calculating the mean and standard errors for monthly precipitation, average monthly maximum temperature, average monthly minimum temperature, and monthly frost days. These were returned to the coop- erators for comment. A relatively .short peri- 'Department of Biology, Montana State University, Bozeman, Montana 59717. 284 I Sept. 1979 Weaver: Fescue Grasslands 285 od of record was used (1) because it seemed wise to compare all stations for the same pe- riod, (2) because many stations lacked a long- er record, and (3) because without contain- ment the number of data points involved would have become unmanageable; 490 data points were collected for each of the 61 sta- tions considered in this study. A grand summary of the climatological data was made by calculating average means and average standard errors across the sta- tions in each type. These data, along with those of the driest and wettest station in each type, are summarized in Figure 1. The driest and wettest station in each type were chosen arbitrarily as those with the lowest and high- est average annual precipitation. The weather stations used were: (1) For Festiica arizonica, Chevlon RS, Flagstaff, Fort Valley, Grand Canyon, Jacob Lake, and McNary, Arizona; Pagosa Springs and Red Feather Lakes, Colorado; and Gascon, Lake Maloya, Los Alamos, Luna RS, Ruidoso, and Wolf Canyon, New Mexico. (2) For Festuca F. ARIZ. DRY F. 40 THURB. WET (c r m) 12 10 8 30 20 ■'V>'- 6 4 10 2 0 - •■Ir 0 -10 -20 --•1 -'I 'f'a'j'a'c ^ (cm) 12 V Fig. 1. Climates of fescue grasslands 1961-1970. The dashed line shows the annual course of mean daily temper- ature; bars reaching above and below it show mean monthly maximum and minimum temperatures; and short bars reaching beyond the cross bars show the mean standard errors of the maximum and minimum. The heavy solid line indicates mean monthly precipitation and heavy bars extending below this line show the standard errors of these means; associated numbers in the F. viridula graphs indicate precipitation levels that lie off the graphs. Arid months (Walter, 1973) are those in which the temperature line rises above the precipitation line. The heavy bar across the base of the graph indicates the number of frost (0 C) days: clear months experience less than one frost per month, hatched months experience from one to six frosts, and solid months experience more than six frosts. 286 Great Basin Naturalist Vol. 39, No. 3 thurberi, Cochetopa Creek, Crested Butte, Silverton, Taylor Park, Telluride, and Wolf Creek Pass, Colorado. (3) For Festuca sca- brella, Babb, Browning, Del Bonito, Elliston, Gibson Dam, Gold Butte, Kalispell, Lewis- ton, Lincoln, Ovando, Phillipsburg, Pole- bridge, Poison, St. Ignatius, and Sula, Mon- tana. (4) For Festuca idahoensis, Bozeman MSU, Gallatin Gateway, Hebgen Dam, Jack- son, Lakeview, Lima, Melville, Mystic Lake, Pony, Red Lodge, Virginia City, White Sul- phur Springs, and Wisdom, Montana; Buffalo 15SW, Burgess Junction, Lamar RS, Wyom- ing, and Kilgore, Idaho. Powder River Pass, Soldier Park, and Willow Park, Wyoming (J. Thilenius); Plummer 3WSW, Moscow, Grangeville, Hill City, Fairfield 8S, and Three Creek, Idaho (M. Hironaka); Dayton, Lacrosse, Pomeroy, Walla Walla, Cheney, Moscow, Pullman, Rosalia, and Goldendale, Washington (Daubenmire 1970); and Bang- tail Ridge, Montana, could probably have been included but were not. (5) For Festuca viridula. Soda Springs, Squaw Valley, Twin Lakes, California; Crater Lake and Santiam Pass, Oregon; Rainier Paradise RS and Ste- vens Point, Washington; and Burke, Idaho. Summer precipitation data for additional sta- tions, usually maintained by the U.S. Forest Service, are recorded in USDC Climatologic- al Data records. Res It was noted earlier that in the southern Rocky Mountains one finds F. arizonica at the lower edge of the forest zone and that in higher forests and meadows one finds F. thur- beri; in the northern Rocky Mountains, F. idahoensis and F. scabrella range from foot- hill grasslands to mountain meadows; west of the Rocky Moimtains one may find F. viri- dula in alpine and subalpine meadows above the normal range of either F. idahoensis or F. scabrella. The following paragraphs, as well as Table 1 and Figure 1, point out similarities and dif- ferences in the climates of the grasslands oc- cupied by these species. The discussion em- phasizes an average climate calculated from measurements made at 6 to 17 weather sta- tions. The average climate for a type will tend to imderestimate precipitation and frost days and to overestimate temperatures be- cause relatively high and inaccessable sites usually lack weather stations. For this reason, the reader should consider carefully the cli- Table 1. Mean climatic characteristic of fescue grasslands. F. arizonica F. scabrella 15 F. idahoensis 17 F. thurberi F. viridula Number of stations 14 6 9 Mean temp., coolest month (C) Daily range (C) Months with less than six frosts -3 17 4 -6 11 3 -8 12 4 -10 20 2 -3 9 4 Mean temp, of coolest frostfree month (C) 13 13 8 12 10 Mean temp., warmest month (C) Daily range (C) 18 17 17 20 17 20 14 19 14 16 Mean SE of maximum temperatures Between years 0.6 0.8 0.7 0.6 0.6 Between stations 0.5 0.3 0.6 1.0 0.8 Mean SE of minimum temperatures Between years 0.5 0.6 0.6 0.5 0.5 Between stations 0.7 0.5 0.6 0.7 0.4 Number of arid months Mean 0 2 0 0 1 Dry year Average annual precipitation (cm) 3 51 3 43 2 48 1 61 2 170 Mean SE of precipitation (mm) Between years Between stations 9 4 7 3 6 4 9 9 25 15 Sept. 1979 Weaver: Fescue Grasslands 287 matic ranges demonstrated by data from ex- treme stands (Fig. 1). Regardless of vegetation type average win- ter air temperature in weather shelters at heights of 1 to 2 m are in the -3 to -10 C range. Night temperatures are 5-10 C cooler than this, and day temperatures are 5-10 C warmer. The daily range is proportional to the distance from the Pacific Ocean, i.e., to continentality. Herbaceous plants and small animals in fescue grasslands usually expe- rience winter temperatures varying only slightly around 0 C because a snow layer in- sulates them from cold air masses. The "frost-free" season is from two to four months long in a fescue grassland: For F. thurberi it is two months, for F. scabrella it is three months; and for the remaining fescues it is from three to four months. Cooler months in the frost-free season have average temperatures of 8-13 C. Warmer months in this period have average temper- atures of 17-18 C in the lower F. arizonica, F. scabrella, and F. idahoensis sites and 14 C in the higher F. thurberi and F. viridula sites. Nightly minimum temperatures average 8-10 cooler and maximum temperatures are 8-10 C warmer than average temperatures. The average standard errors of daily max- imum temperatures taken a month at a time are 0.7 between years and 0.6 between sta- tions. The average standard errors of daily minimum temperatures taken a month at a time are 0.5 between years and 0.6 between stations. The fact that between-year varia- bility exceeds between-station variability is an indication of the homogeneity of temper- ature data gathered in each vegetation type. Temperatures deviating more than two stan- dard errors from the mean are improbable (3 percent). In an average year F. scabrella experiences two arid months, F. viridula experiences one arid month, and the remaining fescues expe- rience no arid months; during these months plant growth is especially dependent on wa- ter stored in the soil profile during the pre- ceeding months. This statement depends on the definition of an arid month as one in which the temperature line, on a graph plotted with the scales used, rises above the precipitation line; this device was developed -by H. Walter (1973) and similar devices are discussed by Daubenmire (1956). Note that if one uses Walter's index the length of arid pe- riods are the same for the driest, the average, and the wettest stand considered in each type, except for F. arizonica (dry) and F. viri- dulii (wet). A "dry year" may be defined as one in which precipitation is always two standard errors below the mean, because the probabil- ity of such low precipitation in one month is about 3 percent. Such a year for the average fescue stand in a type would be drier than the dry stand presented for that type, with the exception of the dry F. arizonica stand. In such a dry year arid months experienced in the frost-free season would be two for F. sca- brella, F. idahoensis, and F. viridula; one for F. arizonica; and none for F. thurberi. Total arid months experienced would be three for F. arizonica and F. scabrella, two for F. ida- hoensis and F. viridula, and one for F. thur- beri. Total precipitation ranges from 43 cm in the F. scabrella type to 177 cm in the F. viri- dula type, but total precipitation is a poor in- dicator of water availability during the grow- ing season. This is shown by two facts: first, the fescue types with the greatest number of arid months (as defined by Walter, 1973) dur- ing an average year include the driest type (F. scabrella) and the wettest type (F. viri- dula), the latter because it receives much of its precipitation during the winter months. Deep, fine-textured soils may compensate for this type of aridity if winter precipitation brings them to field capacity. And, second, within each type tlie wettest stand differs from both the dry and the average stand by the relatively great amounts of precipitation it receives during the winter months. Conclusions A review of climatic data from 61 weather stations representing five fescue grassland types and spanning 25 degrees of latitude leads one to three qualified conclusions. (1) Variation in climate between sites in a fescue grassland type is usually less than variation between years at a site in that type. (2) The climates of fescue grasslands are generally similar, with mean temperatures in the cold- est month between -3 and -10 C, mean tern- 288 Great Basin Naturalist Vol. 39, No. 3 peratures in the warmest month between 14 and 18 C, daily temperature ranges of about 18 degrees, two to four months with fewer than six frost days, and months in which evapotranspiration exceeds precipitation usu- ally less than two. (3) Ecologically important differences between the climates of the fes- cue types lie in summer conditions. Average July temperatures are higher F. arizonica, F. scabrella, and F. idahoensis grasslands (17-18 C) than in F. thurberi and F. viridula grass- lands (14 C). And arid months decline from three in F. arizonica and F. scabrella to 1 or 2 in F. idahoensis, F. thurberi, and F viridida grasslands. A comparison of the fescue climate with that of other vegetation types of the northern Rocky Mountains (Weaver 1979) shows that the fescue grasslands generally appear in a coniferous forest climate: (1) its frost-free sea- son is similar to that of Fseudotsuga trienziesii and Abies lasiocarpa zones, (2) its average July temperatures are similar to those of the Fseudotsuga and Abies zones, (3) its average annual precipitation is similar to that of the Fseudotsuga and Abies zones, and (4) its drought periods are similar to those of the Fseudotsuga and Abies zones. Environmental factors other than temperature and precipi- tation—perhaps wind, snow cover, soil char- acteristics, or fire— must allow fescue grass- lands to dominate the sites they do. Literature Cited Blaisdell, J. 1958. Seasonal development and yield of native plants. USDA Technical Bulletin 1190. Washington, D.C. 68 p. Daubenmire, R. 1956. Climate as a determinate of vege- tation distribution in eastern Washington and northern Idaho. Ecol. Monogr. 26:131-154. 1970. Steppe vegetation of Washington. Wash. Ag. Expt. Sta. Tech. Bulletin 62. Pullman. 1.31 p. Hitchcock, A. 1950. Manual of the grasses of the United States. U.S. Govt. Printing Office, Wash- ington, D.C. 1051 p. KucHLER, A. 1964. Potential natural vegetation of the conterminous United States. Am. Geographic Soc. Special PubUcation .36., N.Y. 1.541 p. MuEGGLER, W. 1971. Weather variations in a mountain grassland in southwestern Montana. USDA For- est Service Research Paper INT 99, Ogden, Utah. 25 p. vanRyswyk, a., a. McLean, and L. Marchand. 1966. The climate, native vegetation, and soils of some grassland at different elevations in British Colum- bia. Can. J. Plant Science 46:35-50. USDC (1961-1970). Climatological data. U.S. Dept. of Commerce, Washington, D.C. Walter, H. 1973. Vegetation of the earth. Springer Ver- lag, N.Y. 237 p. Weaver, T. 1979. Climates of vegetation types of the northern Rocky Mountains and adjacent plains. American Midland Naturalist 101 (In press). STONEFLY (PLECOPTERA) RECORDS FROM THE BASIN RANGES OF NEVADA AND UTAH Andrew L. Sheldon' Abstract.- Distributional records are given for 40 stonefly species on 15 isolated mountain ranges in Nevada and Jtah. The isolated ranges within the Great Basin lave attracted little attention from aquatic biologists, and the running water fauna of the egion is very poorly known. Gather, Stark, md Gaufin (1975) summarized records of Ne- vada stoneflies, and Baumann, Surdick, and Gaufin (1977) cite a few additional records or Nevada and western Utah. The stonefly faunas of regions surrounding the Great Basin are described by Jewett (1959, 1960), Newell and Minshall (1976), and Stewart, Baumann, and Stark (1974). In this paper I document stonefly distribu- tions on 14 mountain ranges in Nevada and the Deep Creek Mountains, Utah, based on repeated collecting trips in 1974 and 1977. These and published data are to be used in a numerical analysis of stonefly distribution within the Great Basin. Records are listed by mountain range and but one record is given for each range. Records redundant with those of Gather et al. (1975) are not listed although I visited many of their localities. Nymphal records are included because the region is so little known and because these data are in- cluded in the quantitative analysis. This research was supported by Grant 7717 from the Penrose Fund of the American Philosophical Society. I thank Dr. R. W. Bau- mann for examining some of the specimens. Nemouridae Malenka sp. A.— Nevada: Clark Co., Spring Mtns., Gold Cr., 2$ S , 3-V-77. This species will be described by R. W. Baumann from these and previously collected speci- mens from the Spring Mtns. I have Malenka nymphs from Deer Cr. and Willow Cr., Spring Mtns. Malenka californica (Claassen).— Nevada: Lander Co., Toiyabe Ra., Birch Cr., ? , prob- ably this species, 24-VI-77; White Pine Co., Schell Creek Ra., Kalamazoo Cr., $ , 2 $ ? , 26-VI-77. Podmosta delicatula (Claassen).— Nevada: Humboldt Co., Santa Rosa Ra., Dutch John Cr., 5 ? ? , 19-VI-77. Prostoia besametsa (Ricker).— Nevada: Elko Co., Jarbidge Mtns., Canyon Cr., ? , 11- VI-74; Ruby Mtns., Thomas Cr., $ , 2 ? ? , 12-VI-74. Zapada cinctipes (Banks).— Nevada: Elko Co., Ruby Mtns., Lamoille Cr., nymphs, 17- 11-77; Esmeralda Co., White Mtns., Chiatov- ich Cr., 4 $ $, ? , 10-11-77; Humboldt Co., Jackson Mtns., Bottle Cr., nymphs, 5-II-77; Pine Forest Ra., Alder Cr., 4 alifor- nia, Nevada, western Utah, Arizona, and southern New Mexico. March to June. 13. Chyptantha Lehm. Catsevc Annual, biennial, or perennial, herbaceous or fruticulose plants; stems erect or ascend- ing, usually with coarse stiff pubescence; leaves opposite at base, or alternate through- out, firm, veinless; flowers white or rarely yellow; inflorescence spikelike or racemose, bracted or bractless; calyx divided to the base, the lobes erect or connivent, linear or oblong, when mature investing the nutlets and falling away entire, or the calyx per- sistent and the nutlets falling away separate- ly; corolla with a short to somewhat elongate cylindrical tube with or without scales at the base of tube, the fornices usually con- spicuous; style slender, short or long, includ- ed; stigma capitate; nutlets 1-4, erect, ovate to triangular, roughened or smooth, winged, margined or marginless, affixed laterally through a median ventral and commonly bas- al forked groove; gynobase usually columnar, subulate, or pyramidal. An exclusively American genus of about 100 species of western North and South America. (From the Greek, cryptos, hidden, and anthos, flower, because of the minute co- rolla in some species.) References Johnston, I. M. The North American Species of Cryptantha. Contr. Gray Herb. 74: 1-114. 1925. Payson, E. B. A Monograph of the Section Oreocarya of Cryptantha. .\nn. Mo. Bot. Card. 14: 211-358. 1927. Higgins, L. C. A Revision of Cryptantha Sub- genus Oreocarya Brigham Young Univ. Sci. Bull., Biol.'Ser. 13(4): 1-63. 1971. 1. Plants annual, with slender stems (of longer duration in C. racemosa) 2 Plants biennial or perennial 34 2(1). Calyx circumscissle at maturity; low diffuse plant; inflorescence compact, each flower in axil of leafy bract 1- C. circumscissa Calyx not circumscissle ^ 314 Great Basin Naturalist Vol. 39, No. 4 3(2). Gynobase subulate, protruding beyond the nutlets, bearing a sessile stigma on its tip; root and base of plant with a purple dye; each flower in the axil of a leafy bract 2. C. micrantha — Gynobase shorter than the nutlets; style developed; root or herbage usually with very little or no dye; flowers all or in part bractless (except C. maritima) 4 4(3). Nutlets roughed or (in C. maritima) at least one of them so 5 — Nutlets smooth and shining, not roughened 31 5(4). Margins of nutlets decidedly winged or knifelike 6 — Margins of the nutlets rounded or obtuse 14 6(5). Pedicels usually evident, slender, 1-4 mm long; lateral angles of nutlets distinctly winged 7 — Pedicels obscure or none, less than 1 mm long 8 7(6). Nutlets homomorphous, broadly winged 3. C. holoptera — Nutlets heteromorphic, narrowly winged 4. C. racemosa 8(6). Lateral margins of the nutlets usually distinctly winged; nutlets 4; calyx symmetrical 9 — Lateral margins of the nutlets knifelike or acute 10 9(8) Corolla conspicuous, 4-7 mm broad; nutlets homomorphic 5. C. oxijgona — Corolla inconspicuous, ca. 1 mm broad; nutlets heteromorphic, the odd one often wingless 6. C. pterocarya 10(8). Nutlets 1 or 2, odd nutlet axial 7. C. utahensis — Nutlets 4; odd nutlet abaxial 11 11(10). Nutlets homomorphous; obscurely roughened 12 — Nutlets heteromorphous, plainly mviricate 13 12(11). Nutlets lucid, somewhat bent, margin thickish 8. C. pnsilla — Nutlets dullish, straight, margin thin, the back high, convex 9. C. costata 13(12). Nutlets 1.3-1.7 mm long, the margins of the lateral angles knifelike; calyx 2.5-3.5 mm long, in fruit; corolla 1 mm broad 10. C. inaequata — Nutlets ca. 1 mm long, the margins of the lateral angles merely sharp; calyx ca. 3 mm long in fruit; corolla 1-2.5 mm broad 11. C. angustifolia 14(5). Nutlets decidedly heteromorphous 15 — Nutlets homomorphic 20 15(14). Mature calyx strongly appressed to the flattened rachis, decidedly gibbous on the axil side, persistent 12. C. dumctorum — Mature calyx somewhat spreading, not at all gibbous 16 16(15). Odd nutlet abaxial, surpassed by style 17 — Odd nutlet axial; style surpassed or occasionally reaching to the nutlet tips 18 17(16). Nutlets 1.3-1.7 mm long; calyx 2-3 mm long 10. C. inaequata — Nutlets ca. 1 mm long, calyx 3-4 mm long 11. C. angustifolia 18(17). Odd nutlet smooth and shiny 13. C. maritima — Odd nutlet tuberculate or pappilate 19 19(18). Spikes bracteate 14. C. minima — Spikes naked 15. C. crassisepala December 1979 Higgins: Boraginaceae of the Southwest 315 20(19). Stvle surpassing the nutlets 21 — Style definitely surpassed by or about reaching to the tips of the nutlets 25 21(20). Spikes bracted throughout 22 — Spikes bractless or onlv sparingly so 2.3 22(21). Plant spring flowering; stems dichotomously branched from the base outward; plant usually low 5-15 cm high, and spreading 16. C. mexicana — Plant summer flowering; main stems straight and erect, forming a central axis producing dichotomously branching laterals; plants usually taller, 15-40 cm high 17. C. albida 23(21). Nutlets bent, lucid, gynobase narrowly pyramidal 8. C. pusilla — Nutlets straight, usually dull, gynobase subulate 24 24(23). Nutlets triangular ovate, with a suggestion of a median dorsal ridge; plant dull dark green 18. C. muricata — Nutlets lanceolate or lance-ovate; plants canescent 19. C. intennedia 25(24). Corolla conspicuous 2-5 mm broad 26 — Corolla inconspicuous 0.5-22 mm broad 27 26(25). Nutlets only 1 or 2 in a normal fruit; style not more than half as long as nutlet . 20. C. decipiens — Nutlets normally 4; style often more than half as long as nutlets 19. C. intermedia 27(25). Nutlets usually solitary 28 — Nutlets usually 4 29 28(27). Mature calyx and nutlet conspicuously recurved or deflexed 21. C. recurvata — Mature calyx and nutlets straight 20. C. decipiens 29(27). Nutlets ovate or triangular ovate 22. C. echinella — Nutlets lanceolate -^0 30(29). Stems spreading hirsute 23. C. harbigera — Stems strigose 24. C. nevadensis 31(4). Spikes bracteate, stems reddish 13. C. maritima — Spikes naked, stems greenish 32 32(31). Style reaching one-fourth-three-fourths height of nutlets; calyx densely h.ispid-villous 25. C. gracilis — Style almost reaching the nutlet tips or surpassing them 33 33(32). Margins of nutlets acute at least above the middle; Californian 26. C. mohavensis — Margin of nutlets rounded or obtuse; plants with a definite central axis; not californian 27. C. fendleri 34(1). Corolla tube elongate, distinctly surpassing the calyx; flowers usually heterostyled 35 — Corolla tube short, scarcely if at all surpassing the calyx; flowers not heterostyled '*'* 35(34). Nutlets smooth and shining 36 — Nutlets more or less roughened or wrinkled at least on the dorsal surface 38 316 Great Basin Naturalist Vol. 39, No. 4 36(35). Corolla yellow .37 — Corolla white 30. C. capitata 37(36). Inflorescence an elongate, cylindrical thyrse; nutlets lanceolate, with acute margins, usually only one developing 28. C. flava — Inflorescence consisting of a large terminal cluster with one or more remote, at maturity frequently stalked, much smaller lateral clusters; nutlets broadly ovate, with winged margins, all four usually maturing 29. C. confertiflora 38(35). Nutlets muricate or vernicose 31. C. fiilvoconescens — Nutlets rugose or tuberculate 39 39(38). Ventral or inner surface of the nutlets smooth or nearly so 40 — Ventral surface of the nutlets distinctly roughened 41 40(39). Corolla tube 7-10 mm long; calyx lobes 5-7 mm long in anthesis; plants not heterostyled; nutlets conspicuously tuberculate and short rugose 32. C. oblata — Corolla tube 12-14 mm long; calyx lobes 7-9 mm long in anthesis; plants strongly heterostyled; nutlets finely tuberculate or rugose 33. C. paysonii 41(39). Inflorescence 0.1-0.4 dm long; corolla tube 10-12 mm long; margins of nutlets not in contact; plants less than 1.2 dm tall 34. C. paradoxa — Inflorescence 0.5-3 dm long; corolla tube 5-10 mm long; margins of nutlets in contact or nearly so; plants usually over 1.2 dm tall 42 42(41). Scar of nutlets surrounded by an elevated margin but tightly closed; style 1-2 mm long; calyx .3.5-4 mm long in anthesis 35. C. hakeri — Scar of nutlets conspicuously open; style 3-8 mm long; calyx 4.5-7 mm long in anthesis 43 43(42). Corolla tube 7-10 mm long; scar of nutlets conspicuously open and surrounded by a definite elevated margin 36. C. flavoculata — Corolla tube 5-7 mm long; scar of nutlets slightly open and with only an inconspicuous elevated margin if any 37. C. tenuis 44(34). Nutlets smooth on their dorsal surface 45 — Nutlets more or less roughened on the dorsal surface 47 45(44). Plants strong perennials; crests at base of corolla tube conspicuous; calyx not noticable accrescent, widespread species 38. C. jajnesii — Plants biennial or weak perennials; crests at base of corolla tube lacking; calyx noticable accrescent 46 46(45). Inflorescence capitate or nearly so; calyx segments in fruit 5-7 mm long, a narrow endemic of Coconino Co., Arizona 39. C. atwoodii — Inflorescence broad topped due to the elongation of the cymules in age; calyx segments in fruit 7-10 mm long; SE New Mexico south into Texas and Mexico . 40. C. pahneri 47(44). Ventral surface of nutlets smooth or nearly so 48 — Ventral surface of nutlets rugose or variously roughened 49 48(47). Nutlets bordered by a conspicuous wing; robust plants 5-10 dm tall, with long ebractate spikes; Arizona 41. C. setosissima — Nutlets never conspicuously winged; plants 2-4 dm tall; inflorescence very broad and bracteate; NE New Mexico 42. C. thyrsifloro 49(47). Scar somewhat constricted below the middle of the open portion, NE Arizona . 43. C. osterhoutii December 1979 HuiGINS: BORAGINACEAE OF THE SoUTHWEST 317 Scar triangular and not constricted below the middle or (closed in C. hakeri) 50 50(49). Cvmules elongating and so the inflorescence broad; biennial or short-lived perennials; nutlets usually with an evident dorsal ridge 51 — Cymules shorter and the inflorescence narrow; long-lived perennials; nutlets with only a slight dorsal ridge if any 53 51(50). Surface of the leaves with inconspicuous appressed bristles; inflorescence open, with only a few elongate cymules, 7-14 cm long terminating the stem; endem- ic to near Las Vegas, Nevada; known only from the type and may not exist any longer due to urbanization in the area 44. C. insolita — Surface of the leaves conspicuously setose-hispid with spreading bristles; inflorescence open, at least at maturity 52 52(51). Calyx lobes 7-12 mm long in fruit; nutlets 3-4.5 mm long, prominately carinate on the dorsal side 45. C. virginensis — Calyx lobes 5-7 mm long in fruit; nutlets 2.5-3 mm long, with only an indistinct central ridge toward the apex 46. C. hoffmannii 53(50). Nutlets indefinitely tuberculate and rugose to nearly smooth; W Nevada and E California 47. C tumulosa — Nutlets definitely tuberculate, rugose or muricate 54 54(53). Scar of nutlets open 55 — Scar of nutlets closed 35. C. hakeri 55(54). Ventral surface of nutlets deeply rugose and tuberculate, the dorsal less so ' 48. C. ahata — Ventral surface of nutlets muricate or verrucose, the dorsal also or with some of the murications connected to form short irregular ridges 49. C. humilis 1. Cryptantha circumcissa (H. & A.) I. M. Johnston Lithospennum circumscifisiim H. & A. Bot. Beechey Voy. 370. 1840. Piptoccihjx circuinscisstts Torr. in S. Wats. Bot. King Exp. 240. 1871. Eritrichium circiiniscissitm A. Gray, Proc. Amer. Acad. Arts 10: 58. 1874. Knjnitzkia circumscissa A. Gray, Proc. Amer. Acad. Arts 20: 275. 1885. Wheele- rella circumscissa Grant, Bull. S. Calif. Acad. Sci. 5: 28. 1906. Greeneocharis circwnscissa Rydb. Bull. Torrey Club .36: 677. 1901. Cryptantha cir- cumscissa I. M. Johnst. Contr. Gray Herb. 68: 55. 1923. {Tolmie, Snake Ft. Snake County, Idaho) Small annual herbs; stems erect or bushy branched, forming round clumps 0.2-1 dm tall, strigose to very hirsute; leaves oblanceol- ate to nearly linear, 0.4-1.5 cm long, 1-2 mm broad, obtuse, strigose or hirsute, pustulate with small inconspicuous pustules, the pe- tioles somewhat siliceous; inflorescence short, congested, the racemes obscure; bracts evi- ■ dent, appearing as if a continuation of the fo- liage leaves; calyx 2.5-4 mm long in fruit, ob- long-ovate, connate to near the middle, the lobes falling away by a circumscission near the sinuses, hirsute, the tube ciliceous, per- sistent; pedicels about 0.5 mm long; corolla minute, white, inconspicuous, 1-2(3) mm broad; style just exceeded by the nutlets or equalling them; gynobase about % height of nutlet; nutlets 4, homomorphous, or with the abaxial one .slightly larger, triangular-ovate or oblong-lanceolate, 1.2-1.7 mm long, mar- gins acute, the surfaces smooth or inconspic- uously muriculate, scar closed and forked be- low. Dry, open, usually .sandy slopes and plains, widely distributed in many plant commu- nities, however in our area found rriostly in the Larrea and Juniperus communities. Cen- tral Wa.shington to Baja California, mostly east of the Cascade and Sierra Nevada ranges to southern Idaho, Utah, and Arizona; also Chile and Argentina. March to July. 2. Cryptantha micrantha (Torr.) I. M. John- ston Eritrichium micranthum Torr. Bot. Mex. Bound. 141. 1859. Knjnitzkia micrantha A. Gray, Proc. Amer. .\cad. Arts 20: 275. 1885. Eremocarya micrantha 318 Great Basin Naturalist Vol. 39, No. 4 E. L. Greene, Pittonia 1: 59. 1887. C. micrantha I. M. Johnst. Contr. Gray Herb 68: 56. 1923. {Thurber, sand hills, Frontera, Texas, and in oth- er places along the Rio Grande, March-April) Eremocarija miiricata Rydb. Bull. Torrey Club 36: 677. 1909. {Parry 164, Valley of the Virgin near St. Gewge) Eritrichiwn micranthum var. lepidum A. Gray. Syn. Fl. N. Amer. 2 pt. 1. 193. 1886. Krynitzkia mi- crantha var. lepida A. Gray, Proc. Amer. Acad. Arts 20: 275. 1885. Eremocarya lepida E. L. Greene, Pittonia 1: 59. 1887. Eremocarya mi- crantha var. lepida Macbr. Proc. Amer. Acad. Arts 51: 545. 1916. Cryptantha micrantha var. le- pida I. M. Johnst. Contr. Gray Herb. 68: 57. 1923. {Cleveland, San Diego, California, 1876) = var. lepida Slender annual herbs; stems dichotomously branched throughout, 0.5-1.5 dm tall, root and lower part of the stem dye stained, stri- gose; leaves linear to oblong-oblanceolate, 0.3-0.7 cm long, 0.8-1.4 mm wide, strigose to villous-hirsute, pustulate on the dorsal side; inflorescence short, dense, 1-4 cm long, the spikes unilateral solitary or geminate, nu- merous; bracts conspicuous, subtending each flower; calyx 1.8-2.5 mm long in fruit, ovate- oblong, slightly asymmetrical, conspicuously biseriate, the segments oblong-lanceolate, hirsute; pedicels 0.5-0.8 mm long; corolla in- conspicuous to evident, 0.5-2.5(3.5) mm broad; style short, the stigma sessile; gyno- hase subulate, much longer than the nutlets; nutlets 4, homomorphous to slightly hetero- morphous, the abaxial one the most persistent and slightly larger, lanceolate with apex at- tenuate, 1-1.3 mm long, margins rounded, plumbeous or brown, smooth or tuberculate, scar extending entire length of nutlet, nar- row, only slightly broadened at the base. Dry, open, sandy slopes and plains. Nevada and Utah, south to Baja California and Ari- zona, eastward to southern New Mexico and Transpecos Texas; also northern Mexico. March to June. The species is easily recognized because of its dense bracteate spikes, dye-stained root, and the long, protruding gynobase. 3. Cryptantha holoptera (A. Gray) MacBride Eritrichiwn holopterum A. Gray, Proc. Amer. Acad. Arts 12: 81. 1876. Krynitzkia holoptera A. Gray, Ibid. 20: 276. 1885. Oreocarya holoptera E. L. Greene, Pittonia 1: 58. 1887. Cryptantha holop- tera Macbride, Contr. Gray Herb. 48: 44. 1916. (£. Palmer, Ehrenber, Arizona, 1876) Coarse annual herbs; stems erect, some- what woody toward the ba.se, 1-6 dm tall, the branches ascending, rather numerous, hir- sute also somewhat strigose; leaves oblanceo- late to linear-lanceolate, 3-6 cm long, 3-8(12) mm wide, hirsute, conspicuously pus- tulate on the dorsal side, less so above; in- florescence racemose, the racemes usually geminate, 0.4-0.7(1) dm long; bracts incon- spicuous or evident on a few racemes; calyx 2.5-3.5 mm long in fruit, oblong-ovate, the segments lanceolate, connivent, midrib thick- ened and hirsute; pedicels ascending or re- curved, 0.7-1.5 mm long; corolla white, min- ute; style evidently surpassing the nutlets; gynobase slender, nearly equalling the nut- lets; nutlets 4, homomorphous, ovate to slightly oblong-ovate, 1.5-2.5 mm long, mar- gins narrowly to broadly winged, the surface of the nutlets dark with lighter tubercula- tions, scar subulate, closed above, clearly open below. Dry, gravelly, or rocky slopes and ridges mostly in the Larrea community. Inyo Coun- ty, California, .south to southern Imperial County, California, and eastward to Mohave and Yuma counties, Arizona, not common. February to April. 4. Cryptantha racemosa (S. Wats.) E. L. Greene Eritrichiwn racemoswn S. Wats, in A. Gray, Proc. .\mer. Acad. Arts 17: 226. 1882. Krynitzkia race- mosa E. L. Greene, Bull. Calif. Acad. Sci. 1: 208. 1855. C. racemosa E. L. Greene, Pittonia 1: 115. 1887. Johnstonella racemosa Brand, Feddes Re- pert. Spec. Nov. Regni Veg. 21: 249. 1925. (S. B. & W. F. Parish 775, canyon near Mesquite Sta- tion, San Bernardino County, California, March 1881) C. suffriiticosa Piper, Proc. Biol. Soc. Wash. 32: 42. 1919. {Orcutt 2070, Camp Muchacho, Colorado Desert) C. racemosa var. lignosa 1. M. Johnst. Univ. Calif. Publ. Bot. 7: 445. 1922. /. racemosa var. lignosa Brand, Feddes Repert. Spec. Nov. Regni Veg. 21: 249. 1925. {Hall & Chandler 7034, Panamint Canyon, Panamint Mountains, California) Long-lived somewhat .suffruticose annual; sterns simple, with many a.scending branches or diffusely branched from near the base, 1-10 dm tall, younger parts green, hirsute and afso strigose, older parts brown, woody, with exfoliating epidermis; leaves oblanceo- late, acute, hirsute, conspicuously pustulate, 1.5-4(6) cm long, 2.5-4(12) mm broad; in- florescence paniculate, the racemes branched and loo.sely flowered, 0.3-1.5 dm long; bracts irregular and inconspicuous; calyx 2-4 mm December 1979 Hl(;C.INS: BORAGINACEAE OF THE SoUTHWEST 319 long in fruit, oblong-ovate, tardily deciduous, the segments linear-lanceolate, strigose and hirsute along the thickened midrib; pedicels 1-4 mm long, slender, frequently recurved; corolla very inconspicuous, about 1 mm broad; style much surpassing the nutlets; gijnohasc subulate, nearly equalling the con- similar nutlets; nutlets 4, heteromorphous, ovate, the acute tips slightlv out-curved, odd nutlet next the abaxial calyx-lobe, 1-2 mm long, subpersistent, muricate or tuberculate or both, consiniilar nutlets 0.8-1.5 mm long, tlie margins narrowly winged, dark with pale tuberculations, scar open or closed above, but opening out into a triangular areola below. Dry, sandy slopes or rocky ridges mostly below 4,500 feet. Inyo County, California, south to northeastern Baja California and eastward to southern Nevada, southwestern Utah, and Mohave and Yuma counties, Ari- zona, not common. March to June. Cryp^uiitha racemosa is the only annual with stems that become somewhat woody or subfruticosc near the base. 5. Cryptantha oxygona (A. Gray) Greene Eritrichium oxygonum A. Gray, Proc. Amer. Acad. Arts 19: 89. 1883. Krynitzkia oxygona A. Gray, Proc. Amer. Acad. Arts 20: 227. 1885. C. oxygona E. L. Greene, Pittonia 1: 120. 1887. (C. G. Pringle, hills bordering the Mohave Desert, Cali- fornia, 1882) Erect annual herbs; stems mostly solitary, 1-4 dm tall, with several well-developed as- cending branches from near the base, strigose also villous-hispid; leaves linear or linear-lan- ceolate, 1-4(6) cm long, 1-2(3) mm broad, strigose or short-hispid, obtusish, pustulate with small numerous inconspicuous pustules; inflorescence dense, the spikes geminate or ternate, 1-3(6) cm long; bracts lacking; cahjx 2.5-4 mm long in fruit, ovate to oblong- ovate, obscurely biserial, the segments lan- ceolate, with somewhat thickened sparsely hirsute midribs, the margins silky strigose, somewhat connivent; pedicels short 0.5 mm long; corolla conspicuous, the limb 4-7 mm broad; style evidently surpassing the nutlets; gynohase about two-thirds as long as nutlets, nearly subulate; nutlets 4, homomorphous, oblong-ovate, 2-2.5(3) mm long, margins nar- rowly winged or knifelike, dorsal side of nut- let low convex, muricate or tuberculate, scar closed or open above, open below with a broadly forked triangular areola. Dry slopes and benches below 5,0(K) feet. California in the inner southern Coastal Ranges from western Merced and Fresno counties to Kern Coimty, western Mohave Desert to Santa Rosa Mountains, Riverside County, eastward just into western Nevada. March to May. A .species closely related to C. pterocarya but having much larger corollas, and ranging more westward, just entering our flora along the western boundary in Kern County. 6. Cryptantha pterocarya (Torr.) E. L. Greene Eritrichium pterocuryttm Torr. Bot. Mcx. Bound. 142. 18.59. Krynitzkia pterocarya A. Gray, Proc. Amer. Acad. Arts 20: 276. 1885. C. pterocarya E. L. Greene, Pittonia 1: 120. 1887. {Pickering & Brackenridge 1047, Walla Walla, Washington; lectotype by I. M. Johnston) Eritrichium pterocaryum var. pectinatum .\. Gray, Proc. Amer. Acad. Arts 10: 61. 1874. Krynitzkia pterocarya var. pectinata A. Gray, Proc. Amer. Acad. Arts 20: 276. 1885. (C. C. Parry 168, 169, in the Virgin River Valley near St. George, Utah) = var. pterocarya. Krynitzkia cycloptera E. L. Greene, Bull. Calif. Acad. Sci. 1: 207. 1884. C. cycloptera E. L. Greene, Pittonia 1: 120. 1887. C. pterocarya var. cycloptera Macbr. Contr. Gray Herb. 48: 44. 1916. (C. G. Pringle, hills near Tucson, .Arizona, 1884) = var. cycloptera Erect annual herb; stems 1-4 din tall, as- cendingly branched from the base and throughout, strigose or very short hirsute; leaves lanceolate to linear, 1-2.5(4) cm long, 1-3(5) mm broad, obtuse, strigose or hispid, dorsal surface conspicuously pustulate, ven- tral surface finely pustulate or the pustules nearly lacking; inflorescence open, the spikes geminate or rarely solitary or ternate, 2-6(12) cm long; bracts inconspicuous or lacking; ca- lyx in fruit (2)3-5 mm long, very accrescent, broadly ovate, the segments ovate to ovate- lanceolate, the midrib slightly thickened and sparsely hirsute, the margins tawny strigose or hirsute; pedicels 0.5-1 mm long; corolla in- conspicuous, 0.5-1(2) mm broad; style sur- passing the body of nutlet but occasionally surpassed by the broad wing margin of nut- let; gynohase slender, about two-thirds height of nutlet; nutlets 4, homomorphous and all winged or heteromorphous and the axial one wingless, the body oblong-lanceolate or lan- ceolate, 2-2.5(3) mm long, margin of nutlet broad and winglike or narrow, entire but usu- ally crenate, entending completely around 320 Great Basin Naturalist Vol. 39, No. 4 the nutlet, surface muricate, scar open or closed above, at the base opening into a di- lated areola. Dry sandy to gravelly washes and bajadas, below 6,000 feet. East of the Cascades and Sierra Nevada from southern Washington to northern Baja California, eastward to south- ern Idaho, Utah, Arizona, and northern So- nora, Mexico. March to June. Cryptantha pterocanja can be separated into two rather weak varieties on the basis of the nutlets as follows: 1. Nutlets heteromorphic, axial one wingless Nutlets homomorphic, all winged var. pterocanja var. cycloptera (Greene) Macbr. Variety pterocanja tends to be more south- erly ranging than variety cycloptera; also there are very few intermediates between the two varieties. 7. Cryptantha utahensis (A. Gray) E. L. Greenell Krynitzkia utahensis A. Gray, Synop. Fl. N. Amer. 2: pt. 1. suppl. 427. 1886. C. utahensis E. L. Greene, Pittonia 1: 120. 1887. Eritrichium ho- lopterum var. suhmoUe .\. Gray, Proc. Anier. Acad. Arts 1.3: 374. 1878. Cniptantha suhmoUis Coville, Contr. U. S. Natl. Herb. 4: 166. 1893. (£. C. Pahner 352. St. George, Utah) Erect ascendingly branched herb; stem solitary at base branched just above the base and throughout, 1-3(4) dm tall, strigose or appressed short hirsute; leaves few, scattered, reduced above, linear to linear-oblanceolate 1-5(7) cm long, 1-4 mm broad, obtu.se, short hirsute, conspicuously pustulate especially on the dorsal surface, less so above; inflorescence open, the spikes geminate or solitary, dense, 0.8-2.5(5) cm long; calyx 2-3(4) mm long in fRiit, ovate to oblong, the ba.se oblique-conic, spreading or recurving, the segments lanceo- late, strongly connivent, the browni.sh thick- ened midrib occasionally bearing spreading or recurved bristles, the margins densely silky villous-hirsute; pedicels obscure; corolla con- spicuous 2-4(5) mm broad; style just slightly shorter than nutlets; gynobase subulate, differing only slightly from the style; nutlets 1 or rarely 2, lanceolate, 1.7-2.5 mm long, margins acute or with a narrow knifelike wing, the .surface pale, muricate, papillate, or occasionally .spinulo.se, the back low convex or flat, scar open, linear and slightly dilated below into a small areola. Dry, sandy or rocky washes and hillsides. Desert region of Inyo, San Bernardino, and Riverside counties, California, eastward through southern Nevada into .southwestern Utah, and western Arizona in Mohave Coun- ty. March to May. 8. Cryptantha pusilla (Torr. & Gray) E. L. Greene Eritrichium pusilluDi Torr. & Gray, Pacif. R. R. Re- port 2; 171. 1856. Krynitzkia pusilla \. Gray, Proc. Amer. Acad. Arts 20: 174. 1885. C. pusilla E. L. Greene, Pittonia 1: 115. 1887. (Pope, Rio Pecos to Llano Estacado, March) Low annual herbs; stems numerous, pros- trate to ascending, very slender, 0.3-1.5 dm tall, canescent, strigose to villous-hirsute; leaves mostly basal, scattered above, linear to linear-spathulate, 1-3 cm long, 1-2 mm wide, hispidulous and pustulate on the dorsal surface, less so ventrally; inflorescence com- pact, the spikes solitary or geminate, 2-8 cm long, densely flowered; bracts lacking or the bracts few and minute; calyx 2-2.5 mm long in fruit, broadly ovate, early deciduous, the segments ovate-lanceolate or oblong- lanceolate, hirsute, the midrib only slightly thickened; pedicels obscure; corolla minute, shorter than the calyx, about 0.6 mm wide; style conspicuously surpassing the nutlets; gynobase narrowly pyramidal, about equal- ling the nutlets; nutlets 4, homomorphous, lu- cid, broadly ovate, bent, 0.8-1.2 mm long, margins acute or knifelike, surface light brown or tan with pale tuberculations, scar subulate and expanded at base into a triangu- lar areola. Dry, sandy or gravelly slopes and washes. Southern Arizona east through southern New Mexico into Trans Pecos Texas; ranging southward into Sonora, Chihuahua, and Du- rango, Mexico. March to May. Cryptantha pusilla is a relatively rare plant which just enters oiu- area along the southern lioundary or the Mexican border. 9. Cryptantha costata Brandegee Cryptantha costata Brandegee, Bot. Gaz. 27: 453. 1899. (Brandegee, Borregos Springs, California, 1895) December 1979 Hi(;gins: Boraginaceae of the Southwest 321 C. seorsa Macbride, Contr. Gray Herb. 48: 46. 19Ifi. (M. E. Jones 3841, Needles, California) Coarse low annual herbs; stems erect, few branched, 1-2 dm tall, densely villous-stri- gose and somewhat hirsute; leaves lanceolate to linear, 1-3 cm long, 2-4 mm wide, dorsal surface hispid, also pustulate, ventral surface villous-strigose and sparsely hispidulous the pustules few and inconspicuous; inflorescence open, the spikes rigid, solitary or geminate, 2-5 cm long; bracts remote, few; calyx in fruit 4-6 mm long, ovate-oblong, deciduous, tlie segments linear lanceolate, connivent with slightly spreading tips, midrib thick- ened, hirsute, margins strigose; pedicels ob- scure; corolla inconspicuous, the tube shorter than the calyx, the lobes ascending; style very similar to the gynobase, much surpassing the nutlets; gynobase subulate; nutlets 4, homo- morphous, or slightly heteromorphous with the nutlet next the abaxial calyx-lobe the largest, triangular or oblong-ovate, 1.6-1.9(2) mm long, margins knifelike or narrowly winged, dorsal surface strongly convex, slightly nigulose or obscurely muriculate, ventral surface flat or slightly concave, scar shallow, closed above opening below into a triangular-subulate areola. Dry sandy washes and bajadas. Inyo Coun- ty to San Diego County, California, eastward just into Arizona in Yuma County. February to May. An interesting plant because of its unusual nutlets which have a flat ventral face and a very high, convex dorsal surface. 10. Cryptantha inaequata I. M. Johnston C. iiuucitiata I. M. Johnst. Univ. Calif. Publ. Bot. 7: 444. 1922. Johnstonella inaequata Brand, Fed- de.s Repert. Spec. Nov. Regni. Veg. 21: 250. 1925. (Hall & Chandler 6925, among rock.s, Pleasant Canvon, Panamint Mountains, Califor- nia, 60() meters altitude) Erect annual herbs; stems ascendingly branched, coarse, 2-4 dm tall, hispid and stri- gose; leaves linear to linear-oblanceolate, 1.5-4 cm long, 1-3(4) mm broad, acute, his- pid, pustulate on the dorsal surface; in- florescence open, the spikes solitary or gemi- nate, 4-12 cm long; bracts few and scattered or lacking; calyx in fruit 2-3(4) mm long, "ovate-oblong, the segments lanceolate midrib moderately thickened and hirsute, axial lobe the most thickened and hirsute; pedicels very short, less than 0.5 mm long; corolla small, 1-2(3) mm broad; style conspicuously sur- passing the nutlets; gynobase narrowly sub- ulate, equalling consimilar nutlets; nutlets 4, heteromorphous, triangular-ovate, margins acute or knifelike, surface brownish with pale tuberculations, odd nutlet about 1.7 mm long, more persistent and slightly lighter in color than the others, next the abaxial calyx- lobe, the consimilar nutlets 1.3-1.5 mm long, .scar subulate, closed above, narrowly triangu- lar below. Dry, ii.sually clay soils, on desert slopes and rocky ridges. Inyo and San Bernardino coun- ties, California, eastward to southern Nevada, southwestern Utah, and western Arizona in Mohave County. March to May. This species is relatively rare throughout its range, but in certain localities, as north- east of Henderson, Nevada, it becomes more common, especially when the moisture sup- ply is sufficient. The species is nearly always on heavy gumbo clay soil. 11. Cryptantha angustifolia (Torr.) E. L. Greene Eiitrichium angustifoliwn Torr. Pacif. R. R. Re- ports 5: 363. 1857. Krynitzkia angustifolia A. Gray, Proc. Amer. Acad. Arts 20: 272. 1885. C. angustifolia E. L. Greene, Pittonia 1: 112. 1887. (Tlioinas, Fort Yuma, Arizona) Diffuse annual herbs; stems much branched from near the base, ascending to nearly decumbent, 0.5-2(3) dm tall, hir.sute to strigose- villous; leaves linear, 1.5-4 mm long, 1-2(4) mm wide, hispid or .strigose, pustulate especially on the dorsal surface; inflorescence rather dense, the spikes geminate, 2.5-6(9) cm long, densely flowered; bracts lacking, ca- lyx in fmit 2-4 mm long, ovate-oblong, as- cending, strongly biseriate, the .segments lin- ear-lanceolate, midrib thickened and hirsute, the margins villous-hirsute and ciliate; pedi- cels obscure, less than 0.5 mm long, corolla inconspicuous to evident, 1-2.5 mm broad; style usually surpa.ssing even the odd nutlet; gynobase columnar, equalling the consimilar nutlets; nutlets usually 4, heteromorphous, ovate-oblong, margins obtuse, acute, or nar- rowly winged, the surface brown with pale tuberculations or murications, odd nutlet next abaxial calyx-lobe, slightly larger than the consimilar nutlets which are about 1 mm long, scar very narrowly linear-lanceolate. Dry, sandy or gravelly washes. South- 322 Great Basin Naturalist Vol. 39, No. 4 eastern California from the Death Valley re- gion to northeastern Baja California and east- ward to southwestern Utah, western Texas, and Sonora, Mexico. March to June. 12. Cryptantha dumetorum E. L. Greene Krynitzkia dumetorum Greene, Pittonia 1: 112. 1887. (Curran, half climbing among bushes at Tehachapi Pass, Cahfornia 1884) Sprawling annual herb; sterns erect, or in age, elongate and scrambling or supported by various shrubs, 1-4(5) dm tall, closely stri- gose; leaves lanceolate, 1-3(4) cm long, 2-4(8) mm wide, thickish, sparsely appressed hispidulous, conspicuously pustulate on the dorsal surface, less so above; inflorescence open, the spikes solitary or geminate, loosely flowered, 5-10 cm long; bracts mostly lack- ing or occasionally with 1 or 2 near the base; calyx in fruit 2-3 mm long, closely appressed to the flattened rhachis, conspicuously asym- metrical, persistent, gibbous at the base on the axial side, the 3 abaxial lobes lanceolate, with thickened hispid midribs, the 2 axial lobes partly united, strigose and deflexed his- pid; pedicels lacking; corolla minute, about 1 mm broad; style subequal to nutlets or slightly shorter than the nutlets; gynohase subulate, narrow; nutlets 4, heteromorphous, ovate-lanceolate, to lanceolate, muricate, odd nutlet axial, persistent, 2-3 mm long, the base enlarged and distorting the calyx, scar open and broad, consimilar nutlets 1.5-2 mm long, deciduous, scar closed or very narrow and linear. Sandy bajadas and hillsides or occasionally in the wash bottoms. Central Mohave Desert of California eastward through southern Ne- vada into southwestern Utah. April to May. The Utah collection of this species is from the west shore of Ivins Reservoir, a consid- erable extension of range from that pre- viously known. The plant probably also oc- curs in Mohave County, Arizona, but has not been documented. 13. Cryptantha maritima E. L. Greene Krynitzkia maritima E. L. Greene, Bull. Calif. Acad. Sci. 1: 204. Aug. 1885. C. maritima E. L. Greene, Pittonia 1: 117. 1887. (£. L. Greene, Guadalupe Island, California, 26 April 188.5) Krynitzkia ramusissima E. L. Greene, Bull. Calif. Acad. Sci. 1: 203. Aug. 1885. non K. ramosis- sima A. Gray 1885. {Mrs. Curran. Mohave Desert, California, 1884) C. maritima var. pilosa I. M. Johnst. Univ. Calif. Publ. Bot. 7: 445. 1922. (Palmer 551, stony ridges, Los Angeles Bay, Lower California) Erect annual herbs; stems reddish, ascen- dingly branched throughout, 1-3(4) dm tall, mostly strigose or occasionally hirsute; leaves linear to lanceolate, acutish, 1-3.5 cm long, 1-3.5 mm wide, sparsely hirsute, coarsely pustulate; inflorescence dense, the spikes soli- tary or geminate 1-7(12) cm long, congested, or glomerate especially when immature; bracts evident, and scattered throughout; ca- lyx in fruit 1-3(3.5) mm long, ovate-oblong, ascending, deciduous at length, the segments linear-lanceolate, connivent, the midrib thickish and hirsute, the margins hirsute-vil- lous to villous; pedicels obscure or lacking; corolla minute, 0.5-1 mm broad; style nearly equalling consimilar nutlet; gynobase sub- ulate one-half-two-thirds length of nutlets; nutlets 1-4, heteromorphous, odd nutlet of- ten the only one developing, abaxial, lanceol- ate, 1-2 mm long, margins rounded, surface smooth and shiny, brownish, scar closed or open at base into a small areola, consimilar nutlets similar, but tuberculate and grayish, early deciduous. Dry washes and desert bajadas. Inyo Coun- ty and throughout southeastern California to northern Baja California and east to southern Nevada, southwestern Utah, Arizona, and So- nora, Mexico. March to May. The variety pilosa I. M. Johnston, is distin- guished from the typical material by the den- sely white-villous calyx-segments. The range of pilosa is scattered within the range of the species. 14. Cryptantha minima Rydb. C. minima Rydb. Vull. Torrey Club 28: 31. 1901. [Rydherg & Vreeland 5697, Cuchara River, above La Veta, Colorado, 2100 m) Small annual herbs; stems erect or ascend- ing-spreading, numerous, 0.5-1.5(2) dm tall, finely strigose and coarselv hirsute; leaves ob- lanceolate, 1-3(4) cm long, 1.5-4 mm broad, obtuse, hirsute or hispid in age, moderately pustulate; inflorescence dense, the spikes soli- tary or occasionally geminate 2-9(15) cm long; bracts evident throughout; calyx in fruit 4-7(9) mm long, oblong-ovate, spreading, asymmetrical, the segments lance-linear, con- nivent, midrib conspicuously thickened and bony, hispid, margins sparsely hirsute or ap- December 1979 HiCCINS: BORAGINACEAE OF THE SoUTHWEST 323 pressed hispid; pedicels short, 0.5-1.5 mm long; corolla small, 1-1.5 mm broad; sti/lc surpassed by odd nutlet, ecjualling or surpass- ing consimilar ones; oynohasc oblong about 0.7 mm long; nutlets 4, hetermorphous, odd nutlet ovate 2-3 mm long, margins angled, the surface brownish, finely muriculate or granulate, persistent, next abaxial calyx-lobe, consimilar nutlets 1.2-1.5 mm long, thick, tu- berculate, scar broadly open especially at the base, not forked. Widely distributed on great variety of soils. Principally on the plains east of the Continental Divide, from Saskatchewan, Canada, south to northern New Mexico and Texas. April to July. This plant is closely related to C. crassise- pala (Torr. & Gray) Greene, a more southern and westwardly growing species. The bracted inflorescences serve best to distinguish it from its southern relative. 15. Cryptantha crassisepala (Torr. & Gray) E. L. Greene EritricJiium crassiscpahun Torr. & Gray, Pacif. R. R. Reports 2; 171. 1857. Krynitzkia cmssisepahi A. Gray, Proc. Amer. Acad! Arts 20: 268. 1885. {Pope, vicinity of Permanent Camp on Rio Pecos, 6-7 April 1856) C. dicarpa A. Nelson, Proc. Biol. Soc. Wash. 16: 30. 1903. (r. D. A. CockereU 30, Mesilla Park, New Mexico) = var. crassisepala C. crassispela var. elachantha I. M. Johnst. Wrightia 2: 20. 1959. (R. McVaugh 8040, north end of Quitman Mountains, 8 miles west of Sierra Blanca, Hudspeth Countv, Texas) = var. elachantha Erect or spreading annual herbs; stems many, ascendingly branched, 0.5-1.5(2.5) dm tall, hirsute to hispid; leaves oblanceolate, 1-3(6) cm long, 2-4(6) mm wide, rounded or obtuse, hirsute, pustulate, the upper scarcely reduced; inflorescence moderately dense, the spikes solitary or geminate 3-10(15) cm long; bracts lacking or 1 to 2 subtending the lower- most flowers; calyx in fruit 5-7(10) mm long, oblong-ovate, slightly asymmetrical, the seg- ments lance-linear, connivent above, midrib very hard and thickened, hispid-hirsute; pedi- cels about 0.5-1 mm long; corolla inconspic- uous to 5 mm in diameter; style surpassed by odd nutlet, equalling or slightly longer than consimilar ones; gynobase narrowly oblong; nutlets 4, or occasionally less by abortion, heteromorphous, odd nutlet persistent, next abaxial calyx-lobe, ovate, acute, 2-2.5(3) mm long, the surface granulate or spinular-muri- cate, brownish, consimilar nutlets earlv de- ciduous, ovate-oblong, 1.2-1.5(2) mm long, tuberculate, scar large, open, occupying most of ventral surface. Usually dry sandy soils on ridges and in washes. Southern Utah and Arizona, eastward to southwestern Colorado, New Mexico, western Texas, and northern Mexico. March to July. 16. Cryptantha mexicana (Brandeg.) I. M. Johnston Krynitzkia mexicana Brandeg. Zoe 5: 182. 1905. C. mexicana I. M. Johnst. Wrightia 2: 161. 1961. (Purpus 8301, near Viesca, southwestern Coa- huila, Mexico, 1903) Dense, low, rounded herbs; stems numer- ous, erect, spreading or ascending, 0.5-2 dm tall, hispid or sparingly strigose-villous; leaves oblong-lanceolate, 2-4(5) cm long, 2-6 mm broad, obtuse, hispid, pustulate, the up- per only slightly reduced; inflorescence dense, very floriferous, the spikes solitary or geminate, 5-15 cm long; bracts evident throughout; calyx in fruit 3-4 mm long, broadly ovate, the segments lanceolate, con- nivent, hirsute to hispid villous; pedicels ob- scure; corolla inconspicuous about 1 mm broad; style barely surpassing nutlets; gyno- base pyramidal, shorter than nutlets; nutlets 4, homomorphous, triangular-ovate, 1-1.3 mm long, margins rounded, the surface tan or brownish with white tuberculations, scar triangular, conspicuously excavated. Exposed slopes and rocky ridges, mostly on limestone or caliche. Southeastern New Mex- ico, western Texas, and southward into north- ern Mexico in the state of Nuevo Leon and Coahuila. March to July. This species is closely allied to C. albida (H.B.K.) Johnston. There should be no con- fusing the two as C. albicki has a straight erect central axis or stem while this species is much branched from the base and through- out; also the flowering times are very differ- ent: C. mexicana is early spring and summer, while that of C. albida is in July and August. 17. Cryptantha albida (H.B.K.) I. M. John- ston Myosotis albida H.B.K. Nov. Gen. et Sp. 3: 91. Aug. 1818. (San Juan del Rio, Queretaro, Mexi- co) Lithospermum ramosum Lehm. Asperif. 2: 328. Nov. or Dec. 1818. Eritrichiiim ramosum DC. 324 Great Basin Naturalist Vol. 39, No. 4 Prodromus 10: 132. 1846. Krynitzkia ramosa A. Gray, Proc. Amer. Acad. Arts 20: 274. 1885. C. ramosa E. L. Greene, Pittonia 1: 115. 1887. (Type probably came from San Juan del Rio, Mexico) Erect annual herb; stems single or more commonly several, with numerous loosely as- cending Ij ranches, the main stem becoming somewhat woody or very stiffened below in age, 1.5-4 dm tall, strigose and sparingly his- pid; leaves spathulate to spathulate-linear, usually folded, 2-3 cm long, 2-5 mm broad, acute to obtuse, dorsal surface hirsute, con- spicuously pustulate, ventral surface sparsely hirsute to nearly glabrous; inflorescence ter- minal on the main stem and the numerous branches, the spikes solitary or rarely gemi- nate, 1-6(10) cm long; bracts numerous, small; calyx in fruit 2.5-3 mm long, ovate, the segments lanceolate, connivent, unequal, hispid; pedicels obscure, nearly sessile; corolla inconspicuous, 1.5-2.5 mm broad; style sur- passing mature nutlets about 0.5 mm; gyno- base pyramidal; nutlets 4, homomorphous, triangular ovate, 1-1.3 mm long, margins rounded, the surface tan or brownish, with low whitish tuberculations, scar triangular, occupying much of ventral face, excavated. Slopes, canyons, and ridges of volcanic or limestone origin. Southeastern Arizona, possi- bly in extreme southern New Mexico. Occur- ring in Trans-Pecos Texas and south in Mexi- co in the states of Sonora, western Coahuila, Chihuahua, Durango, and Queretaro, as well as northwestern Argentina. The species just enters our area in south- eastern Cochise County, Arizona, but may be expected in the Guadalupe Mountains of southern New Mexico. 18. Cryptantha muricata (H. & A.) Nels. & Macbr. Mijosotis inuricata H. & A. Bot. Beecheys Voy. 369. 1840. C. muricata Nels. & Macbr. Bot. Gaz. 61: 42. 1916. {DuHoJas, without locality) Eritrichium muriculatum A. DC. Prodromus 10: 132. 1846. Krynitzkia mtiricuUita A. Gray, Proc. Amer. Acad. Arts 20: 273. 1885. C. muricidata E. L. Greene, Pittonia 1: 113. 1887. (Type not given) C. horridula E. L. Greene, Pittonia 5: .55. 1902. {Mrs. Curran, Salinas River, California, 1885) Krynitzkia denticttlata E. L. Greene, Bull. Calif. Acad. Sci 1: 205. 1885. C. denticulata E. L. Greene, Pittonia 1; 114. 1887. (Curran, western Nevada, 1884) C. dcnsiflora Nels. & Kenn. Proc. Biol. Soc. Wash. 19: 156. 1906. {Kennedy 952, Verdi, Nevada,] 1904) Moderately tall erect annual herbs; stems single or several, 1-10 din tall, ascendingly few to several branched, hirsute also some- what strigose; leaves linear to linear-oblan- ceolate, 1-5(9) cm long, 1-3(4) mm wide, acute, villous-hirsute, inconspicuously pustu- late; inflorescence terminating the main stem and branches, the spikes geminate to quinate, 2-15 cm long; bracts lacking; calyx in fruit 2-4 mm long, ovate, deciduous, the segments lanceolate, very connivent, midrib slightly thickened and tawny-hirsute, the margins his- pid; pedicels obscure; corolla inconspicuous to conspicuous, 1-7 mm broad; style usually much surpassing the nutlets or rarely slightly shorter than them; gynobase linear subulate; nutlets 4, homomorphous, broadly ovate, 1.5-2.5(3) mm long, lucid or dull, muricate or tuberculate, also sometimes granulate, mar- gins acute to rounded, base truncate, scar narrow and nearly closed but at the base broadly forking and with a very small areola. Dry gravelly bajadas and washes, or moun- tain slopes. Southern California from the transition zone to arid desert zone and east- ward to Nevada and Arizona. April to July. Two rather well-defined varieties occur within the area of our flora, the typical varie- ty has conspicuous corollas 2-6 mm broad, while variety denticulata (E. L. Greene) Johnston has inconspicuous corollas 1-2 mm broad; otherwise the plants are quite the same. Two other varieties may occur within our area, variety jonesii (Gray) Johnston, and variety clokeyi (Johnston) Jepson. 19. Cryptantha intermedia (A. Gray) E. L. Greene Eritrichium intermedium A. Gray, Proc. .\mer. Acad. Arts 17: 225. 1882. Krynitzkia intermedia A. Gray, Proc. Amer. Acad. Arts 20: 273. 1885. C. intermedia E. L. Greene, Pittonia 1: 114. 1887. {Nevin, Los Angeles, California 1880-1882) C. quentinensis Macbride. (^ontr. Grav Herb. 56: 58. 1918. {Palmer 608, San Qucntin Bay, Cali- fornia) C. barbigera var. fergusomie Macbr. Ibid. 59. 1918. {Ferguson 42, Palm Springs, California) C. intermedia var. johnstonii Macbr. Ibid. 59. 1918. {Johnston 193S, Claremont. California) Erect annual herbs; stems 1-several, erectly branched, 1.5-5 dm tall, very hirsute December 1979 HiGGINS: BORAGINACEAE OF THE SoUTHWEST 325 with spreading or ascending hairs, also stri- gose; leaves lanceolate to linear, acute to ob- tuse, 2-6(7.5) cm long, 1-5(7) mm wide, hir- sute or strigose, minutely pustulate; itiflorescenee open and lax, the spikes gemi- nate to quinate, mostly ternate, 1-15 cm long; bracts lacking; calyx in fruit (2)4-6 mm long, ovate-oblong, ascending, the segments lance-linear, connivent with spreading tips, midrib moderately thickened and very hir- sute, margins strigose or hispid villous; pedi- cels obscure, 0.5 mm long; corolla con- spicuous, 3-6(8) mm broad; stijle subequal to the nutlets, or slightly longer or shorter than them; gijnobase linear-subulate; nutlets usual- ly 4, or somewhat less by abortion, homo- morphous, lanceolate to ovate, 1.8-2.3 mm long, surface mmicate to tuberculate, grayish or tannish, somewhat granulate also, margins mostly obtuse, scar narrow and linear, or closed but with a small areola at the base. Dry sandy slopes and hillsides. Northern California to northern Baja California mostly west of the Sierra Nevada Mountains but en- tering the desert edge along the eastern foot- hills. March to July. CryptantJia intermedia is a highly variable species and tends to intergrade quite com- pletely with C. barbigera in our area, and to a lesser extent also with C. nevadensis. Tlie larger corolla of C. intermedia will usually separate it from its close relatives. 20. Cryptantha decipiens (M.E. Jones) Heller Knjnitzkki decipiens M. E. Jones Contr. VV. Bot. " 12: 6. 1910. C. decipiens Heller, Miihlenbergia 8: 48. 1912. (M. E. Jones, Yucca, Arizona, 14 May 1884) Slender erect annual herbs; stems ascen- dingly branched throughout, 1-4(5) dm tall, strigose rarely sparsely hirsute; leaves mostly basal, reduced upward, linear, 1-4 cm long, 1-3(4) mm broad, obtuse to acutish, strigose and sparsely hispid, sparsely but evidently pustulate; inflorescence open, the spikes geminate or occasionally ternate or solitary, slender, usually densely flowered, 3-10(14) cm long; bracts lacking; ccdyx in fruit 2-5 mm long, ovate to oblong, strictly ascending, asymmetrical, the segments lance-linear, con- .spicuously connivent with spreading or re- curving tips, the midrib moderately thick- ened and hirsute, margins strigose or weakly hirsute, the abaxial lobe usually slightly the longest; pedicels ob.scure or lacking; corolla minute to evident, 0.8-3.5 mm broad; style short, much surpassed by nutlets; gynobase short pyramidal; nutlet 1 or rarely 2, next abaxial calyx lobe, ovate-lanceolate, 1.5-2.4 mm long, margins rounded, the surface browni.sh, nuiricate to tuberculate, scar nar- rowly linear, but opening at base to form a small areola. Sandy, gravelly, or rocky slopes or hill- sides, often growing on limestone. Inyo and Kern counties, California, south to northern Mexico, and eastward through .southern Ne- vada to Washington county, Utah, and west- ern Arizona. March to May. 21. Cryptantha recurvata Coville C. recurvata Coville, Contr. U. S. Natl. Herb. 4: 165. 1893. {Coville & Funsion 713, Surprise Canyon, Panamint Mountains, California) Sprawling annual herbs; stems slender, as- cendingly branched just above the dye- stained root, 1-3 dm tall, strigose rarely his- pidulose; leaves remote, oblanceolate or lin- ear-oblanceolate 1-2(3.5) cm long, 1-4(5) mm broad, rounded or obtuse, strigose, in- conspicuously pustulate; inflorescence open, the spikes solitary or geminate, slender, 2-10(12) cm long; bracts lacking; calyx in fruit 3-4 mm long, conspicuously asymetric- al, bent and strongly recurved, tardily de- ciduous, the segments linear, the abaxial the longest, midrib moderately thickened and hirsute, the margins appressed hirsute or stri- gose; pedicels lacking; corolla minute, .shorter than the calyx; style much shorter than nut- let; gynobase short and slender; nutlet 1, per- sistent, next abaxial calyx-lobe, lanceolate, in- curved ca. 2 mm long, the tips attenuate- acute, margins obtuse, the surface dull brownish, granulate or muriculate, scar nar- rowly linear or clo.sed above, below opening into a small basal areola. Sandy or occasionally gravelly washes or slopes. Southeastern Oregon south to Inyo county, California, in the Panamint Moun- tains and eastward to San Juan County, Utah, and Mohave County, Arizona. April to June. 22. Cryptantha echinella E. L. Greene C. echinella E. L. Greene, Pittonia 1: 115. 1887. C. andfigua var. echitiella Jepson & Hoover, Fl. Calif. 3: .3.36. 1943. (Sonne, Mount Stanford, above Donner Lake, California 2640 m, 1886) Annual herbs; stems simple below, branched above, with ascending branches, 326 Great Basin Naturalist Vol. 39, No. 4 1-3(4) dm tall, setose or occasionally strigose or hispid; leaves linear to oblance-linear, 1-4(6) cm long, 1-3(4.5) mm broad, obtuse, hispid, pustulate; inflorescence open, the spikes slender, solitary or geminate, 1-5 cm long; bracts evident only near the base; calyx in fruit 4-6 mm long, oblong-ovate, de- ciduous, spreading, the segments linear-lan- ceolate, connivent with spreading tips, midr- ib moderately thickened and hirsute, the margins hispid or strigose; pedicels obscure, about 0.5 mm long; corolla minute, 1-2 mm broad; style slightly surpassed by the nutlets; gynohase narrow, two-thirds as long as nut- lets; nutlets 4, homomorphous, ovoid, 2-2.2 mm long, margins rounded, the surface finely muriculate or granulate, or verrucose, scar very narrowly linear or closed, broadly for- ked at the base. Open diy ridges and slopes in the upper arid transition zone, associated with ]uni- periis, Pintis, and Artemisia. Sierra Nevada Mountains of California eastward to the Charleston Mountains, Nevada, with an iso- lated collection in northwestern Colorado, Moffatt County. Weber and Salamun 12612. Jime to August. 23. Cryptantha barbigera (A. Gray) E. L. Greene EritricJiium harbigerum A. Gray, Synop. Fl. No. Amer. 2; 194. 1878. Krijnitzkia barbigera A. Gray, Proc. Amer. Acad. Arts 20: 273. 1885. C. barbigera E. L. Greene, Pittonia 1: 114. 1887. {Parry 171, Washington County, Utah) Krijnitzkia mixta M. E. Jones, Contr. W. Bot. 13: 6. 1910. (M.E. Jones 5106, St. George, Washington County, Utah) Erect annual herbs; stems 1-several, erectly branched, hirsute, 1-4(5) dm tall; leaves oblong to lance-linear, obtuse, 1-5(7) cm long, 3-7(13) mm wide, hirsute, incon- spicuously pustulate; inflorescence termi- nating the main stem and branches, the .spikes usually geminate, .sometimes solitary or ternate, 2-11(16) cm long; bracts lacking; calyx in fruit 4-8(10) mm long, oblong-lan- ceolate, ascending, the segments lance-linear, with the tips spreading or recurving, midrib moderately thickened and hirsute, the mar- gins long white-villous; pedicels obscure, 0.5-0.8 mm long; corolla inconspicuous, 1-2 mm broad; style subequal to nutlets or slightly longer; gynobase linear; nutlets 1-4, homomorphous, lanceolate, 1.5-2.5 mm long, margins rounded or slightly angled, the sur- face verrucose, brownish, scar linear-lanceol- ate, broadened at the base into a narrowly triangular areola. Dry slopes, wash bottoms, and hillsides. Very common throughout most of the desert southwest from southeastern California and northern Baja California east through south- ern Nevada to southwestern Utah, Arizona, and southern New Mexico into Sonora Mexi- co. February to May. 24. Cryptantha nevadensis Nels. & Kenn. Krijnitzkia barbigera var. inops Brandg. Zoe 5: 228. Sept. 1906. C. nevadensis Nels. & Kenn. Proc. Biol. Soc. Wash. 19: 157. Nov. 1906. C. barbi- gera var. inops Macbr. Proc. Amer. Acad. Arts 51: 548. 1916. (T. Brandegce s.n. Mohave Desert.) C. arenicola Heller, Muhlenbergia 2: 242. Dec. 1906. (Heller 8203, Laws, 3 miles west, Inyo Co., Calif.) C. leptophylla Rydb. Bull. Torrev Club .36: 678. 1909. (Palmer 350, St. George, Utah.) C. nevadensis var. rigida I. M. Johnst. Contr. Gray Herb. 74: 68. 1925. (Pringle, hills bordering the Mohave Desert, California, 1882) Erect or ascending annual herbs; stems 1-several, slender, often flexuous, laxly branched, 1-5 dm tall, closely appressed stri- gose, or rarely sparsely hirsute; leaves linear- oblanceolate to linear, acute to obtuse, 1-4 cm long, 1-5(7) mm broad, sparsely ap- pressed hispid, moderately pustulate; in- florescence lax to somewhat glomerate, spikes geminate or ternate, congested or elongate, 2.5-15 cm long; bracts lacking or occasion- ally bracted at base; calyx in fruit, 4-10(12) mm long, lanceolate, ascending, the segments linear-lanceolate, connivent with slender re- curving tips, midrib tliickened and hirsute, margins villous-setose; pedicels obscure about 0.5 mm long; corolla minute, 1-2 mm broad; style subequal to nutlets or a trifle .shorter; gynobase linear about three-fourths length of nutlets; nutlets 4, homomorphous, lanceolate, 2-2.9 mm long, the margins obtuse, the sur- face mostly verrucose or somewhat muricu- late near the tip, scar narrowly open and lin- ear to nearly closed, but always with a small areola near the base. Dry bajadas, wa.shes, and open hillsides mainly in the Larrea commimity. The stems often .supported by other vegetation. South- eastern California, northern Baja California, and eastward through Nevada to south- December 1979 HuUilNS: BORAGINACEAE OF THE SoUTHWEST 327 western Utah and Arizona. March to May. 25. Cryptantha gracilis Osterh. C. gracilis Osteili. Bull. Torre\- Club .30: 236. 190.3. (Osterhotit 2589, CJlciiwood Spring.s, Garfield Co., Colorado) C. Jiillmanii Nels. & Kenn. Proc. Biol. Soc. Wash. 19: 257. 1906. C. gracilis var. hiUrnannii Munz & Johnst. Bull Torrey Club 49: ,39. 1922. (//i7/- tiuin, Iluffaken Ranch near Keno, Nevada) Slender erect annual herbs; stems 1-sever- al, sparsely branched from the base and above, 1-2(4) dm tall, densely short setose; leaves mostly ba.sal, scattered above, linear- oblong, to narrowly oblanceolate, 1-3 cm long, 1-2(3) mm wide, rounded or obtuse, .se- tose or weakly hispid, incon.spicuously pustu- late; inflorescence open, the spikes solitary or geminate, usually glomerate, 1-2 cm long; bracts lacking; calyx in fruit 2-3 mm long, ovate, spreading, early deciduous, the seg- ments lanceolate, midrib slightly thickened and inconspicuously setose, the margins den- sely setose-villous, often tawny; pedicels lack- ing; corolla minute, less than 1 mm broad; style two-thirds to three-fourths length of nutlet; gynobase about half height of nutlet; nutlets 1 or rarely 2 or 3, homomorphous, lanceolate, 1.5-2 mm long, margins mostly rounded, surface smooth and shiny, scar lin- ear very narrowly open at least at the base. Dry slopes and open areas in the upper Transition Zone. Southern Idaho south through Nevada to Inyo County, California, and east to western Colorado, and northern Arizona. April to July. C. gracilis enters our area from the north, and is found only on some of the higher ranges in the Mohave Desert (Charleston Mountains and the Virgin Mountains). The species never truly grows on the dry desert lowlands. 26. Cryptantha mohavensis E. L. Greene Krynitzkia mohavensis E. L. Greene, Bull. Calif. Acad. Sci. 1: 207. 1885. C. mohavensis E. L. Greene, Pittonia 1; 120. 1887. (Cumin, Mohave Desert, California, 1884) C. fallax E. L. Greene, Pittonia 5: 54. 1902. (£. L. Greene, mountains above Tehachapi, Califor- nia, 22 June 1889) Usually erect annual herbs; stems many branched, 1-4 dm tall, short-hispid to hispid -strigose; leaves linear to linear-lanceolate, 1-4 cm long, 1-3 mm broad, strigose or ap- pressed setose, minutely and densely pustu- late, obtuse; infloresence crowded, the spikes ternate or geminate, usually den.se, 2-6 cm long; bracts lacking; calyx in fruit 3-5 mm long, oblong-ovate, a.scending, deciduous, the segments lanceolate, connivent, midrib mod- erately thickened and often sparsely hirsute, margins commonly silky-strigose; pedicels ob- scure, ca. 0.5 mm long; corolla conspicuous 4-7 mm broad; style evidently surpassing nutlets; gynobase columnar .subulate, three- fourths height of nutlet; nutlets 4, homo- morphous, lance-ovate to lance-oblong, 2-2.5 mm long, margins angled and obtuse near apex, surface smooth and shiny, rarely granu- late, the dorsal side flat or low convex, scar closed above but opening to form a small triangular areola at the base. Dry sandy soils. Southeastern and southern California from Inyo and Kern counties southward to the San Gabriel Mountains and Sierra Libre. May to June. The species just enters our flora along the western boundary of the foothills of the Sierra Nevada near Bishop south to the town of Mohave, California. 27. Cryptantha fendleri (A. Gray) Greene Krynitzkia fendleri A. Grav, Proc. .\nier. .\cad. Arts 20: 268. 1885. C. fendleri Greene, Pittonia 1: 120. 1887. (Fendler, without locality. New Mexico, 1847) C. ramidosissima A. Nels. Erythea 7: 68. 1899. (.\V/- son 5275, Laramie, Wyoming) C. wyomingeasis Gandoger, Bull. Soc-. Bot. Fr. 65: 62. 1918. (Nekon 1523, Cummins, Wyoming) Erect annual herbs; stems solitary with many divaricate or ascending lateral branches, 1-5 dm tall, densely spreading his- pid; leaves narrowly oblanceolate, acute to nearly obtuse, 1-5 cm long, (1)2-4 mm broad, hispid, pustulate on the dorsal surface, much less so above; iyiflorescence broad, the spikes solitary or geminate 2-13 cm long, loosely flowered; bracts lacking or rarely 1 or 2 near the base; calyx in fruit 3-6(7.5) mm long, oblong-lanceolate, a.scending the seg- ments linear to lance-linear, slightly con- nivent with the tips slightly spreading, mid- rib thickened and hirsute, margins strigose; pedicels about 0.5 mm long, obscure; corolla inconspicuous, about 1 mm broad; style sub- equal to or slightly exceeding the nutlets; gijnobase subulate, about two-thirds height of nutlets; nutlets 4, or sometimes fewer by abortion, homomorphous, lanceolate, the tips acuminate, 1.5-2 mm long, margins obtuse or 328 Great Basin Naturalist Vol. 39, No. 4 rounded, surface smooth and usually shiny, scar closed or slightly open above, below forming a triangular areola. Open, exposed, usually sandy sites in the Artemisia and Juniperus associations, 3,500-7,000 feet elevation. Southeastern Washington and northeastern Oregon east to southern Alberta and Saskatchewan to east- ern Nebraska, northern New Mexico, and Arizona. June to August. 28. Cryptantha flava (A. Nels.) Payson Oreocanja flava A. Nels. Bull. Torrey Club 25: 202. 1898. C. flava Payson, Ann. Mo.Bot. Card. 14; 259. 1927. (A. Nelson 3074, point of rocks, Sweetwater County, Wyoming, 1 June 1897) O. hitescens E. L. Greene, Pittonia 4: 93. 1899. C. confertiflora var. hitescens Brand, Pflanzenr. IV. (Heft. 97) 252: 90. 1931. (C. F. Baker, hills about Aztec, New Mexico, 25 .'^pril 1899) Perennial herbs; stems many from a mul- tiple caudex, 1.3-4 dm tall, densely long white-hairy at the base, becoming setose and strigose upward; leaves narrowly oblanceol- ate to nearly linear, acute, 2-9 cm long, 3-8 mm wide, dorsal surface strigose and ap- pressed setose with pustulate hairs, ventral surface almost uniformly strigose, and with the pustules less conspicuous; inflorescence narrow to somewhat open, 0.5-2.5 dm long, conspicuously yellow setose; bracts incon- spicuous; pedicels 3-5 mm long in fruit; calyx 8-10 mm long in an thesis, in fruit becoming 9-12 mm long, the segments linear, densely setose, with yellowish hairs; corolla yellow, the tube 9-12 mm long, crests at base of tube absent or nearly so, fornices yellow, truncate, emarginate, 1-1.5 mm long, limb 8-10 mm broad; style exceeding mature fruit 3-7 mm (heterostyled); nutlets lanceolate, 3.4-4 mm long, 1.9-2.2 mm wide, 1 or 2 usually matur- ing, the margins acute, in contact when more than 1 nutlet matures, both surfaces of nutlet smooth and glossy, scar straight, closed, ele- vated margin lacking. Dry sandy soil often in dune areas, 4,000-7,000 feet elevation. Southwestern Wyoming, south through eastern Utah and western Colorado into northea.stern Arizona and northwestern New Mexico. April to Au- gust. 29. Cryptantha confertiflora (Greene) Pay- son Krynitzkia leiicophaca var. alata M. E. Joites, Proc. Calif. Acad. Sci. 5: 710. 1895. Oreocanja alata k. Nels. Coulter and Nelson, Man. Cent. Rockv Mts. 417. 1909. (M. E. Jones 5144, on sandstone cliffs. Silver Reef, Utah, 3 May 1894) Oreocanja confertiflora E. L. Greene, Pittonia 3: 112. 1896. (S. B. Parish 1316, Cushenberry Springs on the north side of the San Bernardino Moiuitains, San Bernardino County, California, 1882) O. hitea E. L. Greene, Muhlenbergia 2: 240. 1906. name only, Feddes Repert. Spec. Nov. Regni. Veg. 19; 72. 1923. description. (A. A. Heller 8211, White Mountains, Inyo County, Califor- nia, 9 May 1906) Perennial herbs; stems 1-7, slender, 1.7-4.3 dm tall, tomentose at base, strigose and setose upward; leaves linear to oblanceolate, 3-12 cm long, 2-10(16) mm wide, acute, dorsal surface densely strigose and appressed setose with pustulate bases, ventral surface uni- formly strigose and with few or no pustules; inflorescence subcapitate, 0.3-2 dm long, stri- gose, and with flattened, twisted, setose hairs; bracts inconspicuous; calyx in anthesis 6-8 mm long, in fruit becoming 10-14 mm long, the segments linear-lanceolate, strigose and spreading setose; corolla yellow, the tube 9-13 mm long, fornices broad, emarginate, about 1 mm long, crest at base of tube evi- dent or sometimes lacking, limb 8-10 mm wide; heterostyled; nutlets ovate or triangu- lar-ovate, 3.5-4 mm long, 2.5-3 mm wide, usually all four maturing, margins narrowly winged, in contact, both surfaces smooth and glossy, scar straight, closed, and lacking an elevated margin. Dry exposed sites on a wide variety of soil types. Southeastern California, eastward through southern Nevada into northern Ari- zona and southern Utah. April to July. A tall handsome plant closely related to C. flava but having nearly capitate in- florescences and broadly ovate nutlets. The yellow flowers also tend to be lighter in color or a wa.shed out yellow. 30. Cryptantha capitata (Eastw.) I. M. John- ston Oreocanja capitata East, Leaflets W. Bot. 1: 9. 1937. C. capitata I. M. Johnst. J. .\rnold Arbor 21; 66. 1940. (A. Eastwood 5969, Hermit Trail on the south rim of the Grand Canyon, Coco- nino (]()untv, Arizona, 9 .\pril 1917) Erect perennial herbs; stems weak, 1 -sev- eral, 1.5-2.7 dm tall, appressed seto.se; leaves linear or narrowly oblanceolate, 3-8 cm long, 3-5 mm wide, dorsal surface appressed se- tose-pustulate, ventral surface uniformly stri- December 1979 HiGGINS: BORAGINACEAE OF THE SoUTHWEST 329 tj;ose and without pustules; inflorescence capi- tate, or with one or two glomerules below the terminal cluster, 0.1-0.4(7) dm long, spreading white-setose; cahjx 7-9 mm long in anthesis, in fruit becoming 11-16 mm long, the segments linear-lanceolate, conspicuously setose-pustulate; coroUa white, the tube 9-12 mm long, fornices yellow, emarginate, about 1 mm long, papillose, crests at base of tube conspicuous, limb 6-8 mm wide; style ex- ceeding mature fruit 4-5 mm; nutlets lan- ceolate, 4-5 mm long, 2-3 mm wide, two to four usually maturing, the margins in contact, knifelike, both surfaces glossy-smooth, scar closed, straight, and without an elevated mar- gin. Open or exposed somewhat sandy soils in the Transition Zone, 6,500 to 8,500 feet ele- vation. South central Utah and north central Arizona in the Colorado River drainage ba- sin. April to July. In our area this species is restricted to the Grand Canyon National Park along the Kai- bab and Hermit trails, both on the north and south rims. 31. fuh (S. Wats.) Cryptantha ruivocanescens Payson Eritrichiitm glomeratiun var. fulvocanescens S. Wats. Bot. King Exp. 243. 1871. £. fulvocanes- cens A. Gray, Proc. Amer. Acad. Arts 10: 61, 1875. Knjnitzkia fulvocanescens A. Gray, Proc. Amer. Acad. Arts 20: 280. 1885. Oreocanja ful- vocanescens E. L. Greene, Pittonia 1: 58. 1887. C. fulvocanescens Payson, Ann. Mo. Bot. Card. 14: 319. 1927. {Fendler 632, near Santa Fe, San- ta Fe County, New Mexico, 1847) Krynitzkia echinoides M. E. Jones, Proc. Calif. Acad. Sci. 5: 709. 1895. Oreocarya echinoides Macbr. Contr. Gray Herb. 48: 31. 1916. as to synonymy, not as to specimens cited. C. ech- inoides Payson, Ann. Mo. Bot. Gard. 14: .321. 1927. C. fulvocanescens var. echinoides Hig- gins. Great Basin Nat. 29: 30. 1969. (A/. /•;. Jones 5297, Pahria Canvoii, Kane C;()nntv, Utah, 26 May 1894) Densely caespitose perennials from a strongly lignified taproot; stems many from a nuiltiple caudex, 0.8-3 dm tall, white hairy at the base, .setose-hirsute upward; leaves spatu- late or oblanceolate, acute to obtu.se, 1.5-7 cm long, 4-12 mm wide, uniformly strigose, pustules mostly confined to the dorsal sur- face; inflorescence narrow or somewhat open at maturity, 0.3-1.9 dm long, white or yel- lowish setose-hispid; bracts inconspicuous; pedicels 2-10 mm long; calyx 4-6 mm long in anthesi.s, in fruit becoming 9-13 mm long, the .segments linear, densely white or yellow- ish setose-hi.spid; corolla white, the tube 7-11 mm long, fornices yellow, emarginate or rounded, 0.7-1.3 mm long, crests at base of tube evident or lacking, limb 7-9 mm wide; style exceeding mature fruit 3-7 mm; nutlets lance-ovate, 3.5-4.5 mm long, 2-3 mm wide, one or two usually maturing, the margins acute to obtuse, in contact when more than one nutlet matures, both .surfaces densely and uniformly muricate, scar open or nearly closed, elevated margin lacking. Dry, .sandy to clay soils on expo.sed areas in the Artemisia or Juniperus-Pinus association, 4,000 to 7,500 feet elevation. Central Utah and north central Arizona east to western Colorado and central New Mexico, with an isolated population at White Sands National Monument. April to August. Two rather distinct varieties occur within our area and may be separated by the follow- ing key: Murications on the nutlet rounded; corolla 9-13 mm long; inflorescence nar- row, white setose at maturity; usually growing on .sandy soils var. fulvocanescens Murications on the nutlet with one or two setose projections; corolla 7-9 mm long; inflorescence broader and asually yellowish seto.se-hispid at maturity; usually growing on clay soils var. echinoides (Jones) Higgins The variety echinoides is limited in our area to north central Arizona and north- eastern New Mexico. 32. Cryptantha oblata (M. E. Jones) Payson Krynitzkia oblata M. E. Jones, Contr. W. Bot. 13: 4. 1910. Oreocarya oblata Macbr. Proc. .\mer. Acad. Arts 51: 548. 1916. Hemisphaerocarya ob- lata Brand, Feddes Repert. Spec. Nov. Regni Veg. 24: 61. 1927. C. obhta Payson, Ann. Mo. Bot. Gard. 14: 254. 1927. (M. £.' Jones 3579, EI Paso, Texas, 23 April 1884) O. hispidissima Wooton & Standi. Contr. U. S. Natl. Herb. 19: 545. 1915. not O. hispidissima (Torr.) Rydb. {Wright 1566, near El Paso and Dona Ana, March to April) 330 Great Basin Naturalist Vol. 39, No. 4 Perennial or biennial herb; stems several, 1-3.5 dm tall, retrorsely setose and spreading hirsute; leaves oblanceolate, acute, 3-10 cm long, 4-14 mm wide, coarsely strigose and se- tose dorsally with conspicuous pustules, ven- tral surface weakly strigose-setose, and with fewer pustulate hairs, the petioles ciliate- margined; inflorescence somewhat open, es- pecially in age, 0.3-2 dm long, setose-hirsute; calyx 5-7 mm long in anthesis, becoming 8-10 mm long in fruit, the segments linear- lanceolate, densely setose-hirsute; corolla white, tube 7-10 mm long, crests at base of tube lacking, fomices yellow, broad, papil- lose, limb 8-12 mm wide; style 3-5 mm long- er than mature fruit; nutlets ovoid, usually all four maturing, the margins narrowly sepa- rated, acute, 2.5-3 mm long, 2-2.5 mm wide, dorsal surface rugose-tuberculate, ventral sur- face smooth or slightly uneven, scar closed, straight, and without an elevated margin. Sandy or gravelly to rocky hillsides mostly on gypsum soils, 1,000 to 5,000 feet eleva- tion. South central New Mexico south through Trans-Pecos Texas into northern Mexico. March to September. This species is only one of the many gypso- philous plants that occur in the southeastern part of our area. 33. Cryptantha paysonii (Macbr.) I. M. John- ston Oreocarya paysonii Macbr. Contr. Gray Herb. 48: 36. 1916. Hemisphaerocarya paysonii Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 61. 1927. C. paysonii I. M. Johnst. Wrightia 2: 160. 1961. (O. B. Metcalfe 1576, limestone hills at Berendo Creek, Sierra County, New Mexico, 12 May 1905) Caespitose perennials; sterns erect, stout, (0.5)1.6-2.9 dm tall, .strigose and more or less spreading setose; leaves oblanceolate, obtuse to acute, 3-9 cm long, 5-15 mm wide, dorsal surface finely strigose or subtomentose, also setose with pustulate hairs, ventral surface similar but with fewer pustulate hairs; in- florescence subcapitate, consisting of four to six compact cymules, 0.5-1.2 dm long, setose; calyx 7-9 mm long in anthesis, becoming 9-10 mm long in fruit, the segments linear- lanceolate, setose; corolla white to yellowish tinged, the tube 12-14 mm long, crests at base of tube lacking, fornices yellow, rounded, densely papillose, 0.5-1 mm long, limb 10-13 mm wide; heterostyled; nutlets ovate, 2.7-3 mm long, 2-2.5 mm wide, usual- ly all four nutlets maturing, margins narrowly winged, in contact, both surfaces finely rugu- lose or finely tuberculate, scar closed, straight, lacking an elevated margin. Gravelly or rocky hillsides mostly on gyp- sum or limestone soils, 4,000-7,500 feet ele- vation. Southeastern New Mexico and Trans- Pecos Texas in Culberson County. April to June. 34. Cryptantha paradoxa (A. Nels.) Payson Oreocarya paradoxa A. Nels. Bot. Gaz. .56: 69. 1913. C. paradoxa Pavson, Ann. Mo. Bot. Card. 14: .3.30. 1927. (£. P. Walker 91, dry gypsum hills in Paradox Valley, Montrose County, Colo- rado, 17 June 1912) O. gypsophila Payson, Bot. Gaz. 60: 380. 1915. {Payson 458. dry gypsum hills in Paradox Val- ley, Colorado, 18 June 1914) Small perennial herbs; stems 1-several, slender, 0.4-1.2 dm tall, subtomentose near the base, weakly setose above; leaves oblan- ceolate to spatulate, usually folded, obtuse, 1.5-4 cm long, 2-4(7) mm wide, dorsal sur- face with appressed setose-pustulate hairs, ventral surface uniformly strigose and with- out pustulate hairs, the petioles ciliate-mar- gined; inflorescence subcapitate, 0.1-0.4 dm long, setose; bracts inconspicuous; calyx in anthesis 5-6 mm long, in fruit becoming 6-8 mm long, the segments linear-lanceolate, weakly setose; corolla white with a yellow tube 10-12 mm long, crests at base of tube lacking, fornices yellow, broad, slightly emarginate, papillose, 0.5 mm long, limb 10-12 mm wide; stijle exceeding mature fruit 4-9 mm; nutlets lanceolate, turgid, 2-3 mm long, 1.3-1.6 mm wide, all four usually ma- turing, margins acute to obtuse, not in con- tact, dorsal surface densely tuberculate and conspicuously rugose, ventral surface tu- berculate, also somewhat mgulose, scar open, constricted below the middle, the margin ele- vated. Dry, sandy, gravelly, or clay .soils, 4,000 to 7,500 feet elevation. Emery County, Utah, western Colorado, and San Juan County, New Mexico. May to June. In our area known only from one collec- tion by Duane Atwood 2527, 12 miles west of Shiprock on Hwy 504, 15 May 1970. 35. Cryptantha bakeri (E. L. Greene) Payson Oreocarya bakeri E. L. Greene, Pittonia 4: 92. 1899! C. bakeri Payson, Ann. Mo. Bot. Card. 14: 331. 1927. (Baker, Earle, and Tracy 827, Mancos December 1979 HiGGINS: BORAGINACEAE OF THE SoUTHWEST 331 River sa^e plains in southern Colorado, H Jnlv 1898) O. etihphus Rvdh. Bull. Torrey C;iuh 31: (i:37. 1901. (Crandall. Delores, Colorado, 1892) Biennial or short-lived perennials; stems stout, 1-3 dm tall, spreading seto.se-hirsute; leaves oblanceolate, obtuse, mostly basal, 3-6 cm long, 5-12 mm wide, dorsal surface stri- go.se and spreading setose, pustulate, ventral surface uniformly strigose and with few or no pustulate hairs; inflorescence narrow, 0.6-2.5 dm long, setose-hirsute; bracts evident, slightly surpassing the individual cymes; ca- lyx in anthesis 3.5-4 mm long, in fruit be- coming 6-8 mm long, the segments broadly lanceolate or ovate, conspicuously setose; co- rolla white, the tube 4-6 mm long, crests at base of tube lacking, fornices yellow, emargi- nate, 1-1.5 mm long, limb 6-8 mm wide; style exceeding mature fruit 1-2 mm; nutlets ovate-lanceolate, 2.5-3 mm long, 1.5-2 mm wide, three to four usually maturing, margins obtuse, nearly in contact, dorsal surface deeply and sharply rugose, ventral surface tu- bercvilate and short mgose, scar closed, sur- rounded by a definitely elevated white mar- gin. Dry sandy or clay soils in Pinyon-Juniper community 4,000 to 8,000 feet elevation. Southeastern Utah, northeastern Arizona in Apache and Navajo counties, and South- western Colorado. May to August. A species closely allied with C. flavoculata but having a shorter style and corolla, and the nutlet scar tightly closed. 36. Cryptantha flavoculata (A. Nels.) Payson Oreocanjd flavoculata A. Nels. Erythea 7: 66. 1899. C. flavoculata Payson, Ann. Mo. Bot. Card. 14: 334. 1927. (A. Nelson 4572, Piedmont, Wyom- ing, 7 June 1898) O. flavoculata spatulata A. Nels. Erythea 7: 67. 1899. (A. Nehon 2.977, gravelly hilltops near Evanston, Wyoming, 29 May 1897) O. cristata Eastw. Bull. Torrey Club 30: 244. 1903. (Easttvood, Grand Junction, Colorado, 17 May 1893) O. shockleyi Eastw. Bull. Torrey Club 30: 245. 1903. (Shockleij 244, Miller Mountain, Esme- ralda County, Nevada) O. eastwoodae Nels. & Kenn. Muhlenbergia 3: 141. 1908. {Kennedy & Goodding 146, Mormon Mountains, Lincoln County, Nevada) Caespitose perennial herbs; stems 1- several, slender, 1-3.7 dm tall, strigose and spreading setose with slender bristles; leaves linear-oblanceolate to spatulate, obtuse or sometimes acute, 3-11 cm long, 3-15 mm wide, densely strigose and weakly setose, dor- sal surface conspictiously pustulate, ventral surface with few pustules and sometimes silky-strigose; inflorescence narrow, or .some- times slightly open and lax, 0.5-3 dm long; bracts evident but not conspicuous; calyx 5-6 mm long in anthesis, in fruit becoming 8-10 mm long, the .segments lanceolate to ovate; corolla white or pale yellow, the tube usually yellow, 7-10 mm long, crests at base of tube lacking, fornices yellow, minutely papillo.se, 1-2 mm long, limb 8-12 mm wide; style ex- ceeding mature fruit 4-8 mm (heterostyled); nutlets lanceolate to lance-ovate, 2.5-3.5 mm long, 1.8-2 mm wide, usually all four matur- ing, margins obtuse, in contact or slightly separated, dorsal surface muricate, tubercu- late, and with con.spicuous ridges, sometimes nearly foveolate, ventral surface tuberculate, rarely with ridges, scar open, constricted near the middle and surrounded by a high, elevated margin. On a wide variety of soils mostly in the Pinyon-Jimiper community, but also occur- ring in the Artemisia and the Spruce-Fir com- mmiities, 3,000-8,500 feet elevation. East central California eastward through Nevada and Utah into southwestern Wyoming, west- ern Colorado, and northern Arizona. April to July. 37. Cryptantha tenuis (Eastw.) Payson Oreocanja tenuis Eastw. Bull. Torrey Club .30: 244. 1903. C. tenuis Payson, Ann. Mo. Bot. Card. 14: 327. 1927. (A. Eastwood, near Moab, in Court House Wash, Grand Countv, Utah, 25 May 1892) Caespitose perennial herbs; stems slender, 1-many, 1.3-2.5 dm tall, strigose and weakly spreading setose; leaves linear-spatulate, mostly basal, obtuse, 2-5 cm long, 3-6 mm wide, dorsal surface strigose and weakly spreading setose, evidently pustulate, ventral surface uniformly strigose and without pus- tules; inflorescence narrow, interrupted, 0.6-1.4 cm long, weakly setose; bracts incon- spicuous; calyx 4.5-6 mm long in anthesis, in fruit becoming 7-9 mm long, the segments linear-lanceolate, white-setose; corolla white, somewhat campanulate, the tube 5.5-7 mm long, crests at base of tube lacking or some- times evident, fornices yellow, broad, emargi- nate, papillose, limb 5-8 mm wide; style ex- 332 Great Basin Naturalist Vol. 39, No. 4 ceeding mature fruit 3-4 mm; nutlets lanceolate, 3-4 mm long, 1.8-2 mm wide, all four usually maturing, margins acute, nearly in contact, dorsal surface carinate, sharply and deeply rugose, ventral surface rugose, scar open, constricted above the base, and with an elevated margin. Dry, sandy, or clayey exposed slopes and benches, 2,500 to 5,500 feet elevation. South- eastern Utah in Emery, Grand, Wayne, and San Juan counties. The species undoubtedly also occurs in northeastern Arizona, because several collections from San Juan County, Utah, have been made within less than a mile of the Arizona border and may have been within Arizona; it would be very hard to tell exactly where the boundary is in this remote area. April to July. 38. Cryptantha jamesii (Torr.) Payson Eritrichium janicsii Torr. in Marcy, Expl. Red Riv- er 262. 1854. Knjnitzkia jamesii A. Gray, Proc. Anier. Acad. Arts 20: 278. 1885 in part. {James, barren deserts high upon the Platte) E. multicattle Torr. in Marcy, E.xpl. Red River 262. 1854. Oreocanja multicaulis E. L. Greene, Pit- tonia 3: 114. 1896. O. siiffniticosa var. multi- caulis Payson, Univ. Wyo. Pub. Bot. 1: 171. 1926. Hemisphaerocanja suffruticosa var. mtilti- caulis Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 60. 1927. C. jamesii var. multicaulis Payson, Ann. Mo. Bot. Gard. 14: 244. 1927. {Fendler 636, near Santa Fe, New Mexico, 1847) = var. multicaulis. O. abortiva E. L. Greene, Pittonia 3: 114. 1896. Knjnitzkia multicaulis var. abortiva M. E. Jones, Contr. W. Bot. 13: 5. 1910. O. suffruti- cosa var. abortiva Macbr. Proc. Amer. Acad. Arts 51: 547. 1916. Hemisphaerocanja abortiva Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 61. 1927. C. jamesii var. abortiva Payson, Ann. Mo. Bot. Gard. 14: 250. 1927. (S. B. Parish 3694, Bear Valley, San Bernardino Mountains, California, 16-20 June 1895) = var. abortiva. O. cinerea E. L. Greene, Pittonia 3: 113. 1896. O. multicaulis var. cinerea Macbr. Proc. Amer. Acad. Arts 51: 54. 1916. O. suffruticosa var. cinerea Payson, Univ. Wyo. Publ. Bot. 1: 171. 1926. Hemisphaerocarya cinerea Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 61. 1927. C. jamesii var. cinerea Payson, Ann. Mo. Bot. Gard. 14: 246. 1927. (£. L. Greene, southern Colorado, on the plains near Pueblo, 1873) = var. setosa. O. disticha Eastw. Bull. Torrey C:hib .30: 238. 1903. C. jamesii var. disticha Payson, Ann. Mo. Bot. Gard. 14: 248. 1927. (A. Eastwood 90, on Bar- tons Range, San Jvian County, Utah, 13 July 1895) = var. disticha. Knjnitzkia multicatilis var. setosa M. E. Jones, Contr. W. Bot. 13: 4. 1910. Hemisphaerocarya suffruticosa var. setosa Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 60. 1927. C. jamesii var. setosa I. M. Johnst. ex Tidestr. Proc. Biol. Soc. Wash. 48; 42. 1935. {M. E. Jones, near Fo-t Cove, Utah, 27 June 1901) O. pustulosa Rydb. Bull. Torrey Club 40: 480. 1913. C pustulosa Payson, Ann. Mo. Bot. Gard. 14: 2.52. 1927. H. suffruticosa var. pustulosa Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 60. 1927. C. jamesii var. pustulosa Harring- ton, Man. PI. Colorado 466, 641. 19.54. (Ryd- berg & Garrett 9320, Hammond Canyon on the Elk Mountains, San Juan County, Utah, 31 Julv 1911) = VAX . pustulosa O. multicaulis var. laxa Macbr. Contr. Gray Herb. 48: 35. 1916. H. laxa Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 60. 1927. C. jamesii var. laxa Payson, Ann. Mo. Bot. Gard. 14: 246. 1927. (C. G. Pringle 776, on sand hills near Paso del Norte, Chihuahua, Mexico, 20 September 1886) = var. laxa^ Erect to caespitose perennials; stems 1-many, 1-6 dm tall, glabrous to evidently hirsute; leaves linear to broadly oblanceolate, obtuse to acute, 2-15 cm long, 2-15 mm wide, glabrous to hirsute, usually pustulate dorsally, ventral surface lacking pustviles or the pustules very inconspicuous; in- florescence open, the cymules usually elon- gating, tomentose to setose-hirsute; bracts inconspicuous to very evident; calyx in an- thesis 3-4 mm long, in fruit becoming 5-7 mm long, the segments ovate-lanceolate, sub- tomentose to setose-hirsute or sometime nearly glabrous; corolla white, the tube 2.5-3 mm long, crests at base of tube conspicuous, foiTiices light-yellow, emarginate, 0.5-1 mm long, limb 5-8 broad; style exceeding mature fruit 1-3 mm; fruit olDlate-ovoid; nutlets ovate-lanceolate, 1-4 maturing, 2-2.5 mm long, 1.5-2 mm wide, the margins not in con- tact, acute, both surfaces smooth and glossy, scar straight, closed, extending from the base to near the apex, elevated margin lacking. In a wide variety of habitats and on very sandy to extremely gumbo clays, 2,000 to 10,500 feet elevation. Southeastern California eastward through southern Nevada and Utah into Wyoming, South Dakota, southward through the high plains into northern Mexico, also northern Arizona and most of New Mex- ico. April to October. Cryptantha jamesii is a wide-ranging heter- omorphic species with a number of diverse growth forms. These growth forms correlated with soil types and altitudinal differences form the basis for the various varieties. December 1979 Hkxjins: Boraginaceae of the Southwest 333 2(1). 3(2). 4(3). 5(2). 6(5). Ventral surface of the leaves glabrous, the petioles not ciHate niari^ined, or tuf- ted at the base of the plant; in our area limited to northeastern Arizona and northwestern New Mexico var. pustiilo.m (Rydb.) Harrington Ventral surface of leaves strigose or setose, the petioles ciliate margined, leaves usually tufted at the base of plant 2 Stems simple, not branched above the base 3 Stems branched from the base as well as above 5 Stems 1-4.4 dm long, usually twice as long as the basal tuft of leaves; wide- spread variety throughout the higher elevations in Arizona and New Mexico of our area var. miilticaulis (Torr.) Payson Stems 0.2-0.9 dm long, usually not exceeding the basal tuft of leaves 4 Flora bracts exceeding the cymules; stems low, decumbent; mountains of southern California and Nevada var. ahortiva (Greene) Payson Floral bracts not exceeding the cymules; stems erect or nearly so; common on Aiiemisio flats and in the Pinyon-Juniper community, in our area confined to northern Arizona and New Mexico var. setosa (Jones) Johnst. ex Tidestr. Stems decumbent or ascending; plants of the great plains var. jamesii Stems erect 6 Leaves linear; cymules 8 cm long or longer, very lax; in our area confined to sand hills in the vicinity of Las Cruces, New Mexico var. laxa (Macbr.) Payson Leaves oblanceolate, cymules usually shorter than 8 cm long and more con- gested; in our area limited to northern Arizona and northwestern New Mexico on sandy dmie areas var. disticha (Eastw.) Payson 39. Cryptantha atwoodii Higgins C. atwoodii Higgins. Southw. Naturalist 19:(2) 127-130. 1974. (D. Ativood 2624, 7 miles north of Junction Hwy 89/164 on Hwy 89, Coconino County, Arizona, 20 May 1970) Biennial or short-lived perennial herbs; stems several, arising from the branched cau- dex, 0.5-3 dm tall, spreading setose with slen- der somewhat stiffened hairs; leaves oblan- ceolate, folded, obtuse, 1-4 cm long, 2-6 mm wide, setose on both surfaces, conspicuously pustulate on the dorsal side; inflorescence capitate or with several reduced clusters be- low the terminal cymule, 0.1-L3 dm long; calyx 3-4 mm long in anthesis, in fruit be- coming 5-7 mm long, the segments lanceo- late, setose; corolla white, the tube 4-4.5 mm long, crests at base of tube lacking, fornices yellow, rounded, 0.5 mm long, limb 5-8 mm broad; style exceeding mature fruit 1.5-3 mm; fruit depressed globular; nutlets ovate, 1.9-2.5 mm long, 1.8-2 mm wide, usually all -four maturing, margins acute, not in contact, both surfaces smooth and glossy, opaque, scar straight, closed, extending from the base to near the apex, elevated margin lacking. Dry hillsides in shaley soil. A very narrow endemic from Coconino County, Arizona, all collections coming from the area about 7 miles south of the gap along Hwy 89. April to May. The area in which this species grows is ex- tremely overgrazed. It was noted that the sheep in the area also utilized this plant for food, although it is not very palatable; also the individual plants are extremely hard to find. C. atwoodii is one of those rare endem- ics that should be protected. 40. Cryptantha palmeri (A. Gray) Payson Krynitzkia palmeri A. Gray, Proc. Amer. .\cad. Arts 20: 278. 1885. Orcocarya palmeri Greene, Pittonia 1: 57. 1887. Hemisphaerocarya palmeri Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 61. 1927. (Palmer 895, 40 miles south of Sal- tillo, Coahuila, Mexico, March 1880) C. coryi I. M. Johnst. J. Arnold Arbor 20: .396. 1939. (V. L. Cory, s.n., about 2 miles west of Long- fellow, Pecos County, Texas, 15 April 19.36) Biennial or .short-lived perennials; stems 1-several, 1.7-4 dm tall, spreading seto.se or hirsute; leaves linear-lanceolate, acute, 3-10(16) cm long, 4-10 mm wide, strigose and subtomentose, pustulate hairs con- 334 Great Basin Naturalist Vol. 39, No. 4 spicuous on the dorsal surface, fewer and not evident on the ventral surface; inflorescence broad topped due to the elongation of the cy- mules in age, 0.3-2.7 dm long, setose; bracts inconspicuous; calyx 4-6 mm long in an- thesis, in fruit becoming 7-10 mm long, the segments lanceolate, setose or weakly hispid; corolla white, the tube 4-6 mm long, crests at base of tube lacking, fornices yellow, rounded, papillose, 0.5-1 mm long, limb 7-9 mm wide; style exceeding mature fruit 2-3.5 mm; nutlets ovate, 2.5-2.8 mm long, 2-2.7 mm wide, the margins not in contact, acute, both surfaces of the nutlet smooth and glossy, scar tightly closed and without an elevated margin. Gravelly to rock hillsides on gypsum, 1,000-4,000 feet elevation. Southeastern New Mexico, western Texas, and northern Mexico in the states of Nuevo Leon and Coahuila. April to July. A Chihuahuan Desert species that just en- ters our area in southeastern New Mexico. It is found almost exclusively on gypsum or limestone soils. 41. Cryptantha setosissima (A. Gray) Payson Eritrichiwn setosissima A. Gray, Proc. Amer. Acad. Arts 12: 80. 1877. Knjnitzkia setosissima A. Gray, Proc. Amer. Acad. Arts 20: 276. 1885. O. setosissima E. L. Greene, Pittonia 1: 58. 1887. (L. F. Ward 646, at Fish Lake, Sevier County, Utah, 25 August 1875) Biennial or short-lived robust perennial herbs; stems 1-3, erect, 3-10 din tall, hirsute; leaves clustered at the base, reduced upward, oblanceolate, 3-13 cm long, 5-15 mm wide, setose, with some finer twisted pubescence beneath, pustvilate hairs numerous on both surfaces; inflorescence broad topped due to the elongation of the scorpioid racemes, 1-5 dm long; calyx 4-6 mm long in anthesis, in fruit becoming 9-11 mm long, the segments broadly lanceolate or ovate, setose; corolla white, the tube 3-5 mm long, constricted above the ovary by the conspicuous ring of crests, fornices yellow, emarginate, about 0.5 mm long, limb 7-9 mm broad; style exceed- ing mature fruit 1-2 mm; nutlets ovate, 5-6 mm long, 3.5-4.5 mm wide, papery, with a broad winged margin, dorsal surface muri- cate and inconspicuously rugose or tubercu- late, ventral surface smooth or nearly so, scar straight, narrow, slightly open, elevated mar- gin lacking. Gravelly to sandy soils in the Pinyon-Juni- per association or the Spruce-Fir association, 6,000 to 11,000 feet elevation. Nye County, Nevada, eastward to central Utah and south- eastward in the mountainous areas of Arizona to Greenlee County. This is one of the most distinctive species in the entire genus, with its stout, strict, soli- tary stems, and its broadly winged nutlets. 42. Cryptantha thyrsiflora (E. L. Greene) Payson Eritricltitim olomcratum var. Iiispidissimum Torr. Bot. Mex. Bound. 140. 1859 in part. O. hispidis- sima Rydb. Bull. Torrey Club 33: 150. 1906. (Type not given) Oreocanja thyrsiflora E. L. Greene, Pittonia 3: 111. 1896. C. thyrsiflora Payson, Ann. Mo. Bot. Card. 14: 283. 1927. (£. L. Greene, Cheyenne, Wyoming, 6 July 1892) O. iirticacea Wooton & Standi. Contr. U. S. Natl. Herb. 16: 166. 1913. (A. A. & E. G. Heller 3731, Canyoncito, Santa Fe Comity, New Mexico, 18 June 1897) O. monospemia Osterh. Bidl. Torrey Club 46: 55. 1919. (Osterhout 5754, Trinidad, Las Animas County, Colorado, 20 July 1918) Short-lived perennials or sometimes bien- nial; stems stout, 1-several, arising from the base, 1.7-4 dm tall, very hispid; leaves oblan- ceolate, obtuse, 5-12 cm long, 5-14 mm wide, spreading setose or hispid, pustulate on both surfaces; inflorescence very broad 1-3 cm long, 0.6-2.5 dm wide, setose or hispid; bracts 2-3 cm long, but hidden by the elon- gate cymules; calyx in anthesis 3-4 mm long, in fruit becoming 6-9 mm long, the segments linear, setose; corolla white, the tube 3-4 mm long, crests at base of tube conspicuous, for- nices yellow, emarginate, papillose, about 0.5 mm long, limb 5-8 mm wide; style exceeding mature fruit 1-1.5 mm; nutlets ovate to ovate-lanceolate, 2.5-3.5 mm long, 1.5-2 mm wide, usually 2 to 4 maturing, acute, margins in contact, dorsal surface low rugulose and tuberculate, sometimes with murications be- tween the rugae, ventral surface similar but with fewer ridges or sometimes almost smooth, scar subulate, the margin not ele- vated. Plains, foothills, and mountain slopes, 4,500-9,600 feet elevation. Southeastern Wyoming and western Nebraska, south through the eastern two thirds of Colorado into northeastern New Mexico and the Okla- homa Panhandle. May to September. December 1979 HiGGINS: BORAGINACEAE OF THE SoUTHWEST 335 This is a very striking and handsome plant, especially when in full flower. The very broad and rounded inflorescence easily sepa- rates this species from others in the genus. In our area restricted to the northeast quarter of New Mexico. 43. Cryptantha osterhoutii (Payson) Payson OiCiHdnjd o.'itcrhoiitii Payson, Univ. Wyo. Puhl. Bot.'l: 167. 1926. C/ osterhoutii Payson, .'\nn. Mo. Bot. Card. 14: 329. 1927. (G. e'. Ostcrhout 6138, Monument Park, near Grand Junction, Mesa County, Colorado, .3 June 1921) Densely caespitose perennials; stems slen- der, many arising from the densely branched multiple caudex, 0.7-1.2 dm tall, strigose and spreading setose; leaves spatulate to oblan- ceolate, obtuse, 1-3 cm long, 3-8 mm wide, dorsal surface strigo.se and appressed setose, pustulate, ventral surface strigose, the pus- tules mostly lacking; inflorescence open, 0.3-0.8 dm long, weakly white-setose; bracts inconspicu'His; calyx in anthesis 2.5-4 mm long, in fruit becoming 5-6.5 mm long, the segments lanceolate, strigose and spreading setose; corolla white, the tube 2-3 mm long, crests at base of tube usually evident but poorly developed, fornices yellow, broad, emarginate, papillose, about 0.5 mm long, limb 5-7 mm wide; style exceeding mature fruit 0.2-0.7 mm; nutlets lanceolate, 2.7-3.2 mm long, 1.8-2.2 mm wide, usually less than four maturing, margins obtuse, not in con- tact, dorsal surface carinate, sharply tubercu- late and rugose, ventral surface sharply tu- berculate, scar open, con.stricted above the base, elevated margin evident but not con- spicuous. Sandy benches and rocky hillsides, 2,500-6,000 feet elevation. Southeastern Utah, and just into northeastern Arizona and Mesa Coimty, Colorado. May to June. A striking little plant that reaches its great- est concentration in the Canyonlands Nation- al Park area of southeastern Utah. 44. Cryptantha insolita (Macbr.) Payson Oreocarija insolita Macbr. Contr. Gray Herb. 48: 28. 1916. C. insolita Payson, Ann. Mo. Bot. Card. 14: 273. 1927. (L. N. Goodding 2286, Las Vegas, Clark County, Nevada, 4 May 1905) Biennial or short-lived perennial from a slender taproot; stems 1-several, 3-4 dm tall, strigose and abundantly setose; leaves spatu- late, mostly basal, obtuse, 3-5 cm long, 5-14 mm wide, dorsal surface subtomentose and sparsely appressed seto.se pustulate, ventral surface similar but the setae .smaller and few- er, pustules few and inconspicuous, petioles long-hairy at the ba.se; inflorescence open, 0.7-1.4 dm long, cymes few, much elongat- ing, weakly setose; bracts inconspicuous; ca- lyx in anthesis 3.5-4.5 mm long, in fruit be- coming 7-9 mm long, the segments linear lanceolate, densely hirsute; corolla white, the tube 3-4 mm long, crests at base of tube well developed, fornices yellow, slightly emargi- nate, papillose, 0.5-1 mm long, limb 6-8 mm wide; style exceeding mature fruit 1-1.5 mm; nutlets ovate to lanceolate, 3.7-4 mm long, one to four maturing, the margins acute, in contact or nearly so, dorsal surface carinate, tuberculate, granulo-muricate and sometimes slightly rugose, ventral surface tuberculate and somewhat rugulose, scar narrow but open, the margin showing some tendency to become elevated. Alkaline flats and rolling hills, 1,900-2,500 feet elevation. Known only from the region of Las Vegas, Nevada. April to June. A rare endemic that may no longer exist because of the urbanization of the area of Las Vegas. The two known collections were la- beled Las Vegas, so may have occurred in what is now the city or could possibly exist in outlying regions near the town. 45. Cryptantha virginensis (M. E. Jones) Pay- son Knjnitzkia glonierata var. virginensis M. E. Jones, Contr. W. Bot. 13: 5. 1910. Oreocarya virgi- nensis Macbr. Proc. Amer. Acad. Arts 51: 547. 1916. C. virginensis Payson, Ann. Mo. Bot. Card. 14: 274. 1927. (M. E. Jones 5195a, Laver- kin, Washington County, Utah, 8 May 1894) Biennial herbs; stems 1-several, from a stout taproot, 1.5-3(4) dm tall, setose-hirsute with spreading bristles; leaves oblanceolate to spatulate, obtuse, 3-10(12) cm long, 5-15 mm wide, dorsal surface sparsely setose, pu.s- tulate, also with some fine tangled pub- escence beneath, ventral surface sub- tomentose and weakly appressed setose, with only a few pustulate hairs; inflorescence a broad thyrsus with the many individual cymes much elongating, 0.5-3 dm long; bracts conspicuous; calyx in anthesis 3-4 mm long, in fruit becoming 7-11 mm long, the segments linear-lanceolate, hirsute; corolla white, the tube 3-4 mm long, cre.sts at base of tube conspicuous, fornices yellow, emargi- 336 Great Basin Naturalist Vol. 39, No. 4 nate, papillose, about 1 mm long, limb 7-9 mm wide; style exceeding mature fruit 1-1.5 mm; nutlets ovate, 3.3-4.5 mm long, 2.4-2.6 mm broad, usually only one or two nutlets maturing, margins in contact, acute, dorsal surface with a distinct ridge, the surface tu- berculate and usually rugulose, ventral sur- face very uneven with indeterminate rugae and tubercles, scar open and triangular, with an elevated margin. Gravelly to clay soils mostly in the lower sonoran zone, 2,000-8,000 feet elevation. Southeastern California in Inyo and San Ber- nardino coimties, eastward through southern Nevada into Washington County, Utah, and southward into Mohave and Coconino coun- ties of Arizona. March to July. Unlike most of the species of Cryptantho, this showy plant has very fragrant flowers. 46. Cryptantha hoffmannii I. M. Johnst. C. hoffmannii I. M. Johiist. Contr. Arnold Arbor. 3: 90. 1932. Oreocanja hoffmannii Abrams, 111. F!. Pacif. States 3: 600. 1951. (R. Hoffman 78, rocky open slopes of Westguard JPass, Inyo County, California, 11 July 1930) Biennial herbs; stems 1-several, 1.7-3(4) dm tall, conspicuously hirsute; leaves spatu- late, crowded at the base, reduced upward, 2-5 cm long, 5-12 mm wide, spreading se- tose-hirsute, pustulate on both surfaces, but more so dorsally; inflorescence broad topped, interrupted, 1-2.8 dm long; bracts evident but not inconspicuous; calyx in anthesis 3-5 mm long, in fruit becoming 5-8 mm long, the segments lanceolate, hirsute-hispid; corolla white, the tube 3-4 mm long, crests at base of tube evident, fornices yellow, rounded, 0.5 mm long, papillose, limb 5-7 mm wide; style exceeding mature fruit 0.2-0.8 mm; nutlets ovate, 3-3.5 mm long, 2-2.5 mm wide, 2-4 nutlets maturing, the margins in contact, acute, both surfaces irregularly low nigose and minutely tuberculate, the dorsal with a low inconspicuous crest, scar open, triangu- lar, with an elevated margin. Gravelly soils in the Piny on- Juniper associ- ation to the upper transition zone, 7,000-9,000 feet elevation. Southeastern Cal- ifornia in Inyo County and just across the border into Nevada, mostly confined to the area of Westguard Pass. June to July. 48. Cryptantha abata I. M. Johnston Knjnitzkia depressa M. E. Jones, Contr. W. Bot. 13: 5. 1910. not C. depressa A. Nels. Bot. Gaz. 34: 29. 1902. Oreocanja depressa Macbr. Contr. Grav Herb. 48: 32. 1916. C. modesta Pavson, Ann. Mo. Bot. Card. 14: 278. 1927. not C. mod- esta Brand, Feddes Repert. Spec. Nov. Regni. Veg. 24: 48. 1924. C. abata I. M. Johnst. J. Ar- nold Arbor. 24: 240. 1928. (M. £. Jones 6692, Aunun, Nevada, 20 June 1893) Long-lived perennial caespitose herbs; stems many, 0.5-1.8 dm tall, strigose and weakly setose; leaves oblanceolate to spatu- late, obtuse, strigose, setose, and sub- tomentose, the petioles ciliate margined; in- florescence narrow, short, 0.2-0.8 dm long; calyx in anthesis 2.5-4 mm long, in fruit be- coming 5-8 mm long, setose; corolla white, the tube 3-4 mm long, crests at base of tube conspicuous, fornices yellow, rounded, papil- lose, about 0.5 mm long, limb 7-8 mm wide; style exceeding mature fruit 0.5-1 mm; nut- lets in contact, obtuse to acute, dorsal surface carinate, tuberculate, mviricate and some- times with low inconspicuous ridges, ventral surface deeply and irregularly rugose, scar open, triangular, surrounded by a slightly ele- vated margin. Sandy to gravelly soils in the Artemisia and Pinyon-Juniper association, 4,000-9,000 feet elevation. Extreme eastern Nevada, south and western Utah, and Mohave County, Arizona. April to July. Cryptantha abata is a tufted, often mat- forming plant. It is extremely rare in our flora but becomes very common at moderate elevations in Garfield and Piute counties, Utah. 49. Cryptantha humilis (A. Gray) Payson Eritrichium glomcratun) var. humile A. Gray, Proc. Amer. Acad. Arts 10: 61. 1875. Oreocarya hu- milis Payson, Ann. Mo. Bot. Card. 14: 278. 1927. {Bolander, Summit Station, Donner Pass, Nevada County, California, 1871) C. nana var. ovina Payson, .\nn. Mo. Bot. Gard. 14: 314. 1927. C. humilis var. ovina Higgins, Brigham Young Univ. Sci. Bull. 13: no. 4. 37. 1971. (G. H. Benth-y, vicinity of Currant, Nye C'ountv, Nevada, Jiuie 1916) Short-lived perennial herbs; stems many, 0.5-3 dm tall, strigose to spreading setose- hirsute; leaves oblanceolate to spatulate, 1-6 cm long, 2-12 cm wide, strigose, setose, or subtomentose, pustulate on both surfaces; in- florescence narrowly cylindrical to open and lax, 0.2-1.8 cm long, tomentose to con- spicuously setose; bracts inconspicuous; calyx in anthesis 2.5-4.5 mm long, in fruit becom- December 1979 HiCGlNS: BORAGINACEAE OF THE SoUTHWEST 337 ing 6-13 mm long, setose or tomentose; co- rolla white, the tube 2.5-4.5 mm long, crests at base of tube conspicuous to nearly obso- lete, fornices yellow, more or less papillose, rounded, about 0.5 mm long, hmb 7-10 mm broad; style shorter than to exceeding matme fiiiit 2.5 mm; nutlets lanceolate to ovate-lan- ceolate, 3-4.5 mm long, 1.8-3.2 mm wide, 1 to 4 of them maturing, margins in contact, acute to obtuse, dorsal surface muricate, tu- berculate, or somewhat rugulose, ventral sur- face indistinctly muricate or tuberculate, scar open, triangular, margin not elevated. Mostly sandy or gravelly slopes, road cuts, and talus slopes of the higher mountains, 3,500-12,000 feet elevation. Sierra Nevada of California eastward to southeastern Oregon, southern Idaho to western Colorado and ex- treme northwestern Arizona. April to August. Cnjptantha humilis is a common member of the Great Basin flora, but enters our area only in southern Nevada and extreme north- western Arizona. There are 5 varieties in the species complex, with only variety ovina (Payson) Higgins entering our area. 14. Plagiobothrys F. & M. Annual or perennial herbs; stems prostrate to erect, weak to somewhat robust, usually with slender appressed hairs, but at times se- tose though not pungently so; lower leaves opposite, alternate, or rosulate and crowded; flowers borne in slender racemes or spikes, occasionally glomerate, frequently bracted; calyx cleft to near the base, sometimes ac- crescent; corolla white, the tube short and in- cluded in the calyx, the fornices usually prominent and often yellow; stamens includ- ed, the filaments short; nutlets 4, or 1-3 by abortion, erect or incurved, roughened or rarely smooth, tending to be keeled on the back, and with a well-developed ventral keel extending from the tip to the middle or to the base, scar usually elevated and caruncle- like, mostly small, lateral to ba.sal, placed at the base of the ventral keel; gynohase short and broad. About 65 species native to western North America and South America with about 3 outlying species in Australia. (Name from the Greek, plagios, placed sideways, afid bothros, pit or excavation, referring to the position of the nutlet scar.) References Johnston, I. M. A synopsis and redefinition of the genus plagiobothrys. Contr. Gray Herb. 68: 57-80. 1923: and the Allo- carya section of the genus Plagiobothrys in the western U. S. Contr. Arnold Arb. 3: 1-82. 1932. Piper, C. V. A study of Allocarya. Contr. U.S. Nat. Herb. 79-113. 1920. 1. Leaves all alternate, scar lateral, near middle of nutlet 2 Leaves opposite at least below; scar lateral, oblique or basal 9 2(1). Caruncle of nutlet elongate, extending along crest of the ventral keel; nutlets trigonous '^ Canmcle round or nearly so, at or below end of ventral keel 4 3(2). Corolla 4-7 mm broad; nutlets irregularly rugose 1. P.kingii Corolla 1-2.5 mm broad; nutlets conspicuously tessellate 2. P. jonesii 4(2). Canmcle weakly developed, borne at tip of a short or conspicuous stipe; lowest leaves not in a rosette ■'' Canmcle well developed, sessile on the nutlet; lowest leaves mostly in a rosette " 5(4). Stipe of nutlet elongate, about equalling the body in length; nutlets commonly united in pairs, plants of south and west Arizona 3. P. pringlei Stipe of nutlet very short; nutlets distinct; plants mostly Californian 4. P. collinus 338 Great Basin Naturalist Vol. 39, No. 4 6(4). Calyx circumscissle in fruit, less than 4 mm long; lobes usually connivent over fruit; nutlets usually only 1 or 2 7 — Calyx not circumscissle, or, if so, the strongly accrescent calyx over 4 mm long; calyx lobes erect or spreading; nutlets usually 4 8 7(6). Inflorescence a long, simple bracted raceme; nutlets highly incurved in lateral view, 1-2.5 mm long; corolla 2-3 mm broad 5. P. arizonictis — Inflorescence forked, bracted only at base if at all; nutlets low and flattened in lateral view 2-3 mm long; corolla 3-99 mm broad P. nothofuluus 8(6). Transverse dorsal crests of nutlets very narrow and sharp, enclosing polygonal granulate areolas 7. P. canescens — Transverse dorsal crests of nutlets very low and broad, separated only by low lineate ridges 8. P. teneUiis 9(1). Stems strigose or appressed hispidulous 10 — Stems with distinctly spreading hairs; Mohave Desert of California 12. P. parishii 10(9). Scar nearly basal; calyx lobes becoming elongate and thickened, tending all to be directed toward the same side of the fruit; plants mostly prostrate 9. P. leptocladiis — Scar lateral or basilateral, calyx lobes neither elongate nor much thickened, symmetrically disposed; plants prostrate to ascending or erect 11 11(10). Nutlets ovate to lanceolate; the evident scar mostly lateral but occasionally basilateral; plants west of continental divide 10. P. scoideri — Nutlets narrowly lanceolate to lance-linear, scar basilateral, small; plants east of the continental divide 11. P. scopulomm 1. Plagiobothrys kingii (S. Wats.) A. Gray Oregon, extreme eastern California, Nevada, Eritrichium kingii S. Wats. Bot. Kings Exp. 243. and extreme western Utah. May to June. 1871. Plagiobothrys kingii A. Gray, Proc. Amer. Qur plant is variety kinm and iust enters Acad. Arts 20: 281. 188.5. Sonnea kingii E. L. ^u a ■ ..i tvt i \7 ■ . i Greene, Pittonia 1: 2.3. 1887. (S. Wafson 854, ^e flora in southern Nevada. Variety har- eastern side of the Sierra Nevada at Truckee KTiesil (E. L. Greene) Jepson IS a more nor- Pass, California) therly ranging form from northern Nevada Stems erect, 1-several, 1-4 dm tall, hirsute, and California into southeastern Oregon. The also villous-setose; leaves at base of plant nar- <^'y"ies on this phase are more congested and rowly oblanceolate, the cauline lance-linear, usually do not become as elongate. 2-6 cm long, hirsute to hispid, with spread- 2. Plagiobothrys jonesii A. Gray ing or ascending bristles; inflorescence cy- ^'''-;;.i''''/''^^^ ° , ^ . , n "^■5^- l^^f*- Sonnea pnesii E. L. Greene, Pittonia mose, the cymes dense m early flower, SCOr- i, 23. 1887. (M. E. Jones, southeastern California pioid, elongating in fruit and more laxly on the Colorado near the Needles, 5 May 1884) flowered; bracts evident at least on part of Stems erect, simple, widely branched with the inflorescence or flowers; calyx 5-6 mm spreading or ascending branches, 1-3(4) dm long in fruit, the segments lanceolate, very tall, conspicuously hispid and villous-setose, hirsute-hispid; corolla 4-7 mm broad; nutlets the hairs pustulate; leaves oblanceolate to 4, cuneate-ovoid, 2.5-3 mm long, acute and linear at the base 2-6 cm long, the cauline incurved at the apex, dorsal surface with a lanceolate, conspicuou.slv spreading hirsute, low median ridge and similar lateral keels on the hairs with pustulate bases; inflorescence the edges, the whole irregularly rugose with congested when immature, the scorpioid broad papillate areolas; scar elongate, keel- cymes only slightly elongating at maturity like and medial. 1.5-4(7) cm long, some of the lower leaves Dry sandy to gravelly bajadas and valleys also with axillary flowers; bracts lacking; ca- at 4,000-7,000 feet elevation. Southeastern lijx 6-10 mm long in fruit, the segments lin- December 1979 HiGCINS: BORAGINACEAE OF THE SoUTHWEST 339 ear-subulate, spreading hirsute; coroUa 1-2 mm broad; nutlets 2 or 3, incurved, 4 angled bv the dorsal and ventral keels and the lateral ridges, 2.5-3.5 mm long, apex acute, the keel and lateral angles tuberculate, the surface be- tween tessellate; scar narrow, merging into the keel above and with a diverging lateral ridge extending to either side. Gravelly wash bottoms, rocky ridges, and desert bajadas below 5,800 feet elevation. Southern California eastward to southern Ne- vada, southwestern Utah, and western Ari- zona, south into Sonora, Mexico. March to May. Plogiohothrys jonesii differs from all other members of the genus in that it resembles a CryptantJia in habit and an Amsinckia in nut- let characteristics. 3. Plagiobothrys pringlei E. L. Greene Echidiocdiiid arizonicd A. C^ray in Benthani & Hook- ers Gen. PI. 2: 854. 1876; Proc. Anier. Acad. Arts 11: 89. 1876. non P. arizonicus (A. Gray) Greene. Phigiobotluys pringlei E. L. Greene, Pittonia 1: 21. 1887. {Dr. Smart, Verde Mesa, Arizona) Stems several to many, branched from near the base, prostrate or decumbent to nearly erect, slender, 1-4 dm long, spreading setose with fine short hairs; leaves numerous below, gradually reduced above, narrowly oblan- ceolate to linear, 2-4(6) cm long, 2-5 mm broad, obtuse to acute at apex, appressed stri- gose or canescent to conspicuously seto.se; in- florescence an elongate .spike, floriferous to near the base of the .stem; bracts conspicuous, 1-2 cm long; c«/i/.t 3-4.5 mm long in fruit, the .segments linear-lanceolate, canescent; co- roUa 2-3 mm broad, inconspicuous; nutlets 4, those near the base of stem commonly joined in pairs, the upper separate, ovate, acute at apex, 1.8-2 mm long, dorsal keel evident near the apex but fading to distinct tubercu- lations below, the surface also rugulose with short ridges; scar elevated on a prominent stipe at least 1.3 mm long, and usually as long as the nutlet. On .sandy or gravelly desert flats and ba- jadas. Common in Cochise, Maricopa, Pima, and Pinal counties of Arizona, and northern Sonora, Mexico. March to April. 4. Plagiobothrys collinus (Ph.) I. M. Johnston Plagiobothri/s ccilifornicus var. fiilcescens I. M. Johnst. Gontr. Gray Herb. 68: 74. 1923. AUoca- ryastrum ursimim var. fuhescens Brand, Pflan- zenr. IV 252: 101. 1931. Echidiocarya californica suhsp. fuhescens L. Abranis. 111. Fl. Pacific- States 3: 571. 1951. P. collinus var. fuhescens Higgins Great Basin Nat. .^4(2): 165. 1974. (7". .S. Brandegee, Santa Barbara, Galifornia, 1881) P micrunthus A. Nels. Anier. J. Bot. 2.5: 115. 1938. (A. Nelson 10232. Prescott, .Xri/.ona, moist creek banks, 28 April 1925) Stems .slender, elongate, prostrate or de- cumbent, 1-4 dm long, hispidulous; leaves oblanceolate, obtusish to acutish, 1-3 cm long, 3-5 mm broad, hirsute; inflorescence an elongate spike, remotely flowered and very slender; bracts lacking above the middle of inflorescence; calyx 2.8-3.2 mm long in fruit, the .segments linear-lanceolate, hispidulous; corolla 2-2.5 mm broad; nutlets 4, ovoid, 1.5 mm long, dorsal keel thin above, reduced to a mere line and fading out about middle of nutlet, irregularly nigose, also muriculate; scar on a short stipe near base of nutlet. Dry, open flats, mesas, and valleys, ascend- ing to moderate elevations in the foothills. Southern California, northern Baja Califor- nia, Mexico, eastward to westward Arizona and Sonora, Mexico, and in Chile of South America. Febniary to May. P. collinus is divided into five rather di.s- tinct varieties in western North and South America. The only phase that enters our area is variety fulvescens, with its elongate laxly flowered spikes and harsher, more pene- trating pubescence. 5. Plagiobothrys arizonicus (A. Gray) E. L. Greene Eritrichium canescens var. arizonicwn .\. Gray. Proc. Amer. Acad. Arts 17: 227. 1882. Plagiobothrys arizonicus E. L. Greene ex A. Gray, Proc. .\mer. Acad. Arts 20: 284. 1885. (C. G. Pringle 364, near Camp Lowell, Arizona, 9 .\pril 1881) Stems loosely ascending to erect, usually branched below the middle, 1-4(5) dm tall, hirsute-hispid, also somewhat villous, the ba.s- al part of the stem and the root highly charged with a purple dye; leaves linear-ob- lanceolate, 1.5-5(6) cm long, 2-6(10) nun broad, hirsute, with pustulate hairs, the mid- vein and its branches strong dye stained; in- florescence spikelike, elongate, and remotely flowered, 3-15 cm long; bracts mostly lack- ing on all flowers but with several scattered along the spikes; calyx 3-3.5 mm long in fruit, lobed to about the middle, circum- scissle, the segments connivent and narrowly lanceolate, hirsute and somewhat villous; co- rolla 2-2.5 mm broad; nutlets 1-4, commonly 340 Great Basin Naturalist Vol. 39, No. 4 2, ovoid, abruptly acute at apex, the dorsal surface with rectangular smooth areolae marked off by narrow tuberculate ridges and rugae; scar median in a sunken area at base of keel. Dry desert slopes and mesas, often near the base of rocky outcrops, extending to moder- ate elevations in the mountains 7,000 feet. Western edge of the San Joaquin Valley, Cal- ifornia, eastward through southern Nevada to southern Utah, New Mexico, and south into Sonora, Mexico. March to May. 6. Plagiobothrys nothofulvus (A. Gray) A. Gray Eritrichium nothofulvum A. Gray, Proc. Amer. Acad. Arts 17: 227. 1882. Plagiobothrys nothofulvus A. Gray, Proc. Amer. Acad. Arts 20: 28.5. 188.5. {Douglas, California) Stems 1-several, simple or more often as- cendingly branched from the base, 1.5-5(6) dm tall, villous-hispidulous with spreading hairs, base of plant often slightly dye stained; leaves at base oblanceolate 3-10 cm long, 5-20 mm broad, acute at apex, sparsely vil- lous-setose, cauline leaves few, linear- lanceolate to lanceolate; inflorescence elon- gate, loosely flowered, racemes often paired, 5-15(20) cm long; bracts lacking; calyx 2-3 mm long in fruit, lobed to about the middle, circumscissle, the segments narrowly lanceol- ate, fulvous-hirsute; corolla 6-9 mm broad, showy; nutlets 1 to 4, 2-3 mm long, round- ovoid, abruptly constricted to an acute apex, loosely rugulose-reticulate and somewhat granular tuberculate; scar annular, median at the base of the narrow ventral keel. Open grassy slopes, fields, and roadsides, mostly below 2,500 feet elevation. Southern Washington along the Columbia River, south through Oregon and California on the west slope of the Sierra Nevadas to the Coastal Ranges to northern Baja California, Mexico, occasionally at the desert edge in eastern Kern County, California. March to May. Plagiobothrys nothofulvus just enters our flora along the extreme western boundary in California. 7. Plagiobothrys canescens Benth. Plagiobothrys canesceiis Benth. pi. Hartweg. .326. 1849. Eritrichium canescens A. Gray, Proc. Amer. Acad. Arts 10: 57. 1874. (Hartweg, Sacra- mento Valley, California) P. microcarpa E. L. Greene, Pittonia 1: 21. 1887. (Mrs. R. M. Austin, Butte County, California, May 1883) P. canescens var. apertus E. L. Greene, Pittonia 1: 21. 1887. (£.L. Greene, plains of the upper San Joaquin, 1884) Stems many, branched from the base, de- cumbent or prostrate, rarely erect, 1-4(6) din long, villous or finely hispidulous; leaves lin- ear to linear-oblanceolate, 1.5-5 cm long, 2-7 mm broad, the cauline well developed; inflorescence elongate and loosely flowered in age, 5-25 cm long; bracts conspicuous and well developed, 1-2 cm long; calyx in fruit 4-6 mm long, the segments lanceolate, dense- ly iTifous-villous-tomentose; corolla 2.7-3.5 mm broad; nutlets mostly 4, round-ovoid, abruptly constricted to the narrow acute apex, strongly incurved, obscurely tubercu- late, but with conspicuous transverse rugae forming rectangular papillate intervals; scar median, annular, slightly raised. Gravelly to clayey slopes, plains, and grassy hillsides, also alkaline flats, mostly be- low 4,500 feet elevation. Nearly throughout the length of California, mostly west of the Sierra Nevada, entering the Mohave Desert in Inyo, Kern, and San Bernardino counties. March to May. 8. Plagiobothrys tenellus (Nutt.) A. Gray Myosotis tenella Nutt. ex Hook. J. Bot. Kew Gard. xMisc. .3: 295. 1851. Eritrichium tencllum \. Gray, Proc. Amer. Acad. Arts 10: ,57. 1874. (Gei/eV, "mountains along the Coeur daleine River," Idaho) Stems 1-several, slender, erect or ascend- ing, 1-3 dm tall, .soft-villous; leaves mostly basal, rosettelike, lance-oblong to lance-ellip- tic, 1-4 cm long, 2-8 mm broad, sessile, caul- ine leaves few, ovate to lanceolate, shorter than the basal ones; inflorescence open, loos- ely flowered, tending to elongate in age, slen- der, 4-15 cm long; bracts evident only near the base; calyx in fruit 3-5 mm long, the seg- ments ovate-lanceolate, short villous, whitish or fulvous; corolla 2-3 mm broad; nutlets usually 4, 1.5-2.5 mm long, thick cniciform, u.sually light colored, sharply ridged dorsally and on the edges, tuberculate on the ridges, smooth and shiny between the ridges; scar .small, set just below middle of nutlet at end of keel. Gras.sy, sandy, or gravelly slopes, hillsides, and dry open areas below 5,000 feet eleva- tion. Common from California to British Co- lumbia and Idaho, becoming rather rare in Utah and Nevada, and with several highly December 1979 Hku >: BORAGINACEAE OF THE Soi TMWKST 341 scattered locations in Graham, Gila, Mari- copa, Pinal, and Pima counties of Arizona. March to June. 9. Plagiobothrys leptocladus (E. L. Greene) I. M. Johnston Eritrichiiini ((ilifoniiiuni \ar. siilx^loilnilidltnu A. Gray, Bot. Calif. 1: 526. 1876. Knjnitzkia Califor- nia var. suhglodiidiata A. Gray, Proc. Amer. .\cad. .\rts 20: 266. 1885. AUocanja californica var. suhglociiidiata Jepsoii, Fl. W. Middle Calif. 44.3. 1901. AUocarija suhglochidiata Piper, Contr. U. S. Natl. Herb. 11: 485. 1906. (S. Watson H51. Clover Mountains, Elko Count v, Nevada, l.ecto- type by Johnston) Allocarya leptoclada E. L. Greene, Pittonia .3: 109. 1896. Plaoiohothnjs leptocladus I. M. Johnst. Contr. Arnold Arbor. 3: 38. 1932. (E. L. Crccnc. Pine Creek, Eureka Coiuitv, Nevada, 20 [ulv 1896) Stems prostrate, 1-3(7) dm long, somewhat succulent, sparsely strigose to subglabrous; leaves linear or linear-oblanceolate, 3-8 cm long, 2-5 mm broad, 1 or more pair near the base opposite, dorsal surface sparsely stri- gose-pustulate, subglabrous above; in- florescence spikelike, elongate, loosely flow- ered to near base of plant, the spikes somewhat unilateral; bracts evident at least below; calyx very accrescent, in fruit becom- ing 4-8 mm long, the segments linear, slightlv thickened and succulent, all tending to be directed toward the same side of the ftiiit; corolla minute, 1-2 mm broad; nutlets 1-4, lanceolate, 1.5-2.5 mm long, dorsal sur- face rugo.se-tuberculate, granulate, or penicil- late-hairy, ventral surface angulate, keeled the entire length; scar basal or nearly so, not surrounded by a ridge. Moist depressions of clay flats, lusually in alkaline soils. Oregon south to northern Baja California, Mexico, eastward to western Wyoming and northern Utah, entering the desert edge in Kern and San Bernardino counties of California. April to July. In northern Utah P. leptocladus often forms prostrate mats a meter or more in di- ameter from a single plant, but this is rela- tively rare throughout most of its range. 10. Plagiobothrys scouleri (H. & A.) I. M. Johnston Allocarya cusickii E. L. (ireene. Pittonia 1: 17. 1887. ria^iohothnis cusickii I. M. Johnst. C^ontr. Arnold Arbor. .3: 63. 1932. (W. C. Cusick. Union County, Oregon, in 1883) = var. cusiitiialli A. A. Heller. Bull. Torrev Club 26: 550. 1899. (MacDou^al .95, near Mor- mon Lake, south of Flagstaff, .Xrizona, 12 June 1898) Stems ascending, l-.several, 0.8-2.5 dm tall, glabrous; leaves at base oblong-oval to obovate, petiolate, 2-5 cm long, 1-2.8(4) cm broad, glabrous, pustulate, the cauline leaves, sessile, oblong-lanceolate to ovate, 2-4 cm long, 0.5-2 cm broad; inflorescence a modi- fied dense scorpioid cyme, not much elongat- ing in age; bracts lacking; calyx in anthesis 5-6 mm long, in fruit becoming 7-10 mm long, divided to below the middle, the seg- ments lance-oblong, ciliate; pedicels 1-10 mm long, glabrous; corolla tube 8-9 mm long, glabrous within, the limb 5-6 mm broad, moderately expanded; fornices con- spicuous, glabrous; anthers 2.5-3 mm long, subequal to the filaments; style usually ex- ceeding the corolla; nutlets rugose, inner sur- face slightly concave, the margin forming a collar. Moist rich soil at medium elevations, 6,000-9,000 feet. Coconino and Yavapai coimties, Arizona. 16. Eritrichi Schrad. Depressed-pulvinate perennial plants; stems 2-10 cm tall, or .sometimes acaules- cent; leaves small, usually densely hairy, crowded on the nmnerous .short shoots or a base of the elongate stem; inflorescence a false raceme or spike terminating the short stem, naked or leafy bracteate; pedicels erect; calyx cleft nearly to the base; corolla blue, rarely white, often with a yellow eye, salver- form, with a short, narrow tube; fornices well developed; filaments attached well down in the corolla tube; anthers included; ovary 4 lobed; stigma 1; nutlets 1-4, smooth, at- tached basilaterally to the low stout gyno- base, the apex obliquely truncate, this por- tion surrounded by an entire or toothed margin. A genus of about 4 species of Eurasia and western North America. (From the Greek erion, wool and trichos, hair, referring to the wooly pubescence of E. nanuni, the original .species.) Reference Wight, William R. The genus Eritricliium in North America. Bull. Torrev Club 29: 407-14. 1902. 1. Eritrichium nanum (Vill.) Schrad. Miiosotis luinii X'ill. Prosp. 21. 1779. liiitrichiiini uci- niini Schrad. .^sperif. 16. 1820. Omphaloclcs nana A, C;ray, Proc. Amer. Acad. Arts 20: 26.3. 1885. Lappiila nana Car. in Pari. & Car. Fl. Ital. 6: 861. 1886. (Presumably from the Alps) Eritrichium aretioides var. elongatum Rvdb. Mem. N.Y. Bot. Card. 1: 327. 1900. E.' elongatum Wight, Bull. Torrey Club. 29: 408. 1902. E. na- num var. elongatum Cronq. Vase. PI. Pacif. N. W. 4: 203. 1959. {Rijdherg & Besseij 4891, Span- ish Basin, Montana, 26 June 897) = var. elonga- tum. Eritrichium argcutcum Wight, Bull. Torrey (;iub. 29: 411. 1902. E. elongatum var. argenteum 1. M. Johnst. Contr. Gray Herb. 70: 53. 1924. E. na- num ssp. villosum var. villo.sum i. argenteum Brand, Pflanzenr. IV 252 (Heft. 97): 191. 1931. (Crandall & Coican 361, northwest of Como, Colorado, 31 July 1895) = var. elongatum. Pulvinate-caespitose, long-lived perennials; stems acaule.scent or caulescent with short, slender, erect stems, 0.1-0.7(1) dm tall, vil- lous to densely strigose; leaves oblanceolate to oblong or narrowly ovate, 5-10 mm long, 1-2(3) mm broad, villous to loo.sely .strigo.se; inflorescence compact when sessile among the leaves or racemelike when borne on a leafy flowering branch, capitate; calyx 1.8-2.3 mm long in fruit, linear, villous or silky strigose; corolla tube short, 2-2.5 mm long, yellowi.sh, the limb blue rarely white 4-8 mm broad; fornices prominent, papillo.se; nutlets 1-4, glabrous, somewhat asynunetric- al, margined, with an entire or toothed mar- gin. Open rocky slopes, dry meadows, and on tundra at high elevations in the mountains, 10,000-14,000 feet elevation. Irregularly from the Alps of Europe, acro-ss Asia to 346 Great Basin Naturalist Vol. 39, No. 4 Alaska and south in the Rocky Mountains to northern New Mexico. June to August. Eritrichium as here considered is a highly variable and widespread circumboreal spe- cies, with several varieties. In our flora only variety elongatwn occurs and is limited to only the highest mountain peaks in northern New Mexico. 17. Lappula Gilib. Stickseed Annual or biennial herbs; stems ascending or erect; leaves alternate, entire, narrow, firm, and veinless; inflorescence terminal, the flowers borne in a sympodial, branched cyme; calyx 5-parted, nearly to the base, ac- crescent; pedicels usually erect, short; corolla blue or white, rather inconspicuous, more or less funnelform, with conspicuous fomices; stamens included; variously inserted; style in- cluded; nutlets 4, ovoid to oblong, trigonous or flattened, with 1-3 rows of cylindrical, conical or flattened spines or glochidia on the sides, or on the cupulate border, attached to the elongate gynobase only part of their length. A genus of about 10 species of wide distri- bution in the northern hemisphere (dimin- utive of the Latin lappa, a bur.) Reference Johnston, I. M. Studies in the Boraginaceae. A synopsis of the American native and immigrant borages of the subfamily Bo- raginoidae. Contr. Gray Herb. 70: 47-51. 1924. 1. Nutlets with 2 rows of slender marginal prickles that are not confluent at base; corolla about 3 mm broad , 1. L. echinata Nutlets with a single row of marginal prickles that are more or less confluent at the base; corolla 2 mm or less broad 2. L. redowskii 1. Lappula echinata Gilib Mijosotis lappula L. Sp. PI. 131. 1753. Lappula myosotis Moench. Meth. 417. 1794. EcJiino'ipci- mum lappula Lehm. Asperif. 121. 1818. Lappula lappula Karst. Deuts. Fl. 979. 1882. {Europe) Lappula echinata Gilib. Fl. Lithu. 1: 25. 1781. {Eu- rope) Echinospermum fremontii Torr. Pacif. R. R. Reports 12: 46. 1860. Lappula fremontii E. L. Greene, Pittonia 4: 96. 1899. {Freinont 844, Pass Creek, near southern end of the Sierra Nevada) Stems simple to freely branched, 1.5-8 dm tall, villous-hirsute; leaves linear to linear- lanceolate or lanceolate, acute or obtuse, nar- rowed to a sessile base, 2-5 cm long, 2-7 mm broad, hispidulous; calyx 2.5-3(4) mm long in fruit, the segments linear, appressed hispi- dulous; pedicels 1-3 mm long, erect; corolla bright blue, 2-4 mm broad; nutlets 3-4 mm long, sharply verrucose or muricate dorsally, with 2 marginal rows of long, slender bristles that are distinct to near the base, these some- times irregularly distributed over the back. Dry plains, hillsides, roadsides and waste places, also cultivated ground. Native to Eu- rasia, but widespread as a weed in northern United States and Canada. June to August. L. echinata is rare in our flora, known only from Schultz Pass, Coconino County, Ari- zona (W/jj7/ng ii73B). 2. Lappula redowskii (Hornem.) E. L. Greene Myosotis redoivskii Hornem. Hort. Bot. Hafn. 1: 174. 1813. Echinospermum redoivskii Lehm. .Vsperif. 127. 1818. (Russia) Echinospermum texanum Scheele, Linnea 25: 260. 1852. Lappula texana Britt. Mem. Torrey Club. 5; 273. 1894. L. redoicskii var. texana Brand, Pflanzenr. IV 252 (Heft. 97): 150. 1931. {Roemer, San Antonio, Te.xas) = var. cupulata Echinospermum redowskii var. occidentale S. Wats. Bot. King E.\p. 246. 1871. Lappula redowskii var. occidentale Rydb. Contr, U. S. Natl. Herb. 3: 170. 1895. L. occidentalis E. L. Greene, Pittonia 4: 97. 1899. E. occidentale K. Schum. Just. Bot. Jahresb. 27; 522. 1901. (S. Watson 86L from the Sierra's to the Wasatch) = var. redowskii Echinospermum redowskii var. cupulatum \. Gray, Bot. Calif. 1: 5.30. 1876. Lappula cupulata Rvdb. Bull. Torrey Club 28: 31. 1901. Echinospermum cupulatum K. Schum. Just. Bot. Jahresb. 29: 564. 1903. L. redowskii var. cupulata M. E. Jones, Bull. Univ. Mont. Biol. 15: 44. 1910. (S. Watson 862, Trinity Mountains, Nevada) Lappula desertorum E. L. Greene, Pittonia 4: 95. 1899. Echinospermum desertorum K. Schum. Just. Bot. Jahresb. 27: 522. 1901. L. redowskii var. desertorum I. M. Johnst. Contr. Arnold Arb.r 3: 93. 1932. (£. L. Greene, near Holborn, Nevada, 16 July 1896) December 1979 HlGGINS: BORAGINACEAE OF THE SoUTHWEST 347 Liipptila heterospcnnd E. L. C^ieene, Pittonia 4: 94. 1899. L. tcxana var. Iictewspenna Nels. & Machr. Bot. Gaz. 61: 41. 1916. [Baker. Earh\ 6c Tmcij 826, from near Mancos, in sonthwesltMii Colo- rado) = var. ciipulata Lappula cownata E. L. (Greene, Pittonia 4: 94. 1899. EcluiiosjX'iDiiini rowiKitum K. Schvmi. Just. Bot. Jahresb. 27: 522. 1901. L. tcxaiui var. cownata Nels. & Macbr. Bot. Gaz. 61: 41. 1916. (C. C. Piingle, mesas near Tucson, Arizona. 18 April 1884) = var. cupulata L. leucotricha Rydb. Bull. Torrey Club. 36: 676. 1909. (Touiuct/, near Tucson, Arizona, 20 April 1894) Stems usually simple, or with several uiinor steuis arising from the base of the major stem, or bushy branched from the base, 1-5 dm tall, cinereous hispid-villous; leaves narrowly oblanceolate to spathulate, the basal ones 1.5-5(8) cm long, 3-8 mm broad, rosettelike, the cauline leaves gradually reduced in size upward; inflorescence cymose, the individual racemes terminating the stems and branches; bracts conspicuous, subtending each flower; calyx in fruit 3-5 mm long, the segments lin- ear or linear-lanceolate, strigose; pedicels erect or ascending 1-3 mm long; corolla blue or whitish, 1-2 mm broad; nutlets 2-3 mm long, muricate dorsally, with a single row of nearly distinct prickles, or sometimes with a greatly swollen cupulate border. A weed in dry, usually disturbed areas along roadsides, abandoned fields and waste places. Eurasia and western North America. March to July. Lappula redowskii is a widespread and var- iable species. Many names have been placed on the various forms that occur throughout the range of the species. The most distinct of these forms has been called L. texana, and, if it weren't for the many intermediate charac- teristics between it and the typical L. re- dowskii, it could easily be maintained as a distinct species. It is the many named and nameless forms that occur between these two extremes that have prodticed the abundant .svnonvmv. 18. Hackelia Opiz. Stickseed Ascending or erect biennial or perennial herbs; leaves alternate, broad and veiny; flowers in naked or only basally bracteate scorpioid cymes paniculately disposed; calyx cut to the base into spreading ovate to ob- long or lanceolate lobes; pedicels slender, re- curving in fruit; corolla white or blue, with a short or elongated tube, and an evidently 5-lobed limb, the lobes rounded and connate less than one-third their length; fornices well developed; stamens included, affixed at middle of tube; filaments slender, short; an- thers oblong to elliptic; style slender, scarcely if at all surpassing the nutlets; stigma capi- tate; nutlets 4, erect, ovate to lanceolate, at- tached ventrally to the pyramidal gynobase by a broad medial or submedial areola, the margin with subulate glochidiate prickles which are frequently confluent at the base, the back smooth or with glochidiate appen- dages. A genus containing about 45 species, cen- tering in western North America with out- lying species in South America and Eurasia. References Gentry, J. L. 1974. Studies in the genus Hack- elia (Boraginaceae) in the western United States and Mexico. South- western Nat. 19:139-146. Gentry, J. L. and R. L. Carr. 1976. A revision of the genus Hackelia (Boraginaceae). Mem. New York Bot. Gard. vol. 26, no. 1. 2(i; 3(2). Corolla Hmb white to ochroleucous to greenish tinged 1. //■ ursina Corolla limb blue or occa.sionally violet blue or pink 2 Intramarginal prickles absent on all nutlets (or rarely present on //. florihunda and then only on less than half the nutlets) ^ Intermarginal prickles present on all nutlets 4. H. pinetorum Corolla limb inconspicuous, only 1.5-2.5 mm wide, calyx segments ca. 1 mm long '. 2. H. besseyi 348 Great Basin Naturalist Vol. 39, No. 4 - Corolla limb broader, mostly 4-8 mm wide; calyx segments mostly 1.5 mm long or more 4 4(3). Cymes conspicuously bracteate throughout; cauline leaves long ciliate; stems generally stiffly hirsute 3. H. hirsuta - Cymes bracteate, if at all, only at the base; cauline leaves not long ciliate; stems with appressed hairs 5 5(4). Fornices curved inward at the tips, about twice as long as broad; principal marginal prickles of all mature nutlets less than 2 mm long; inflorescence open and spreading, the branches few; plants slender 4. H. pinetorwn Fornices relatively straight, not curving inward at the tip, about as broad as long; principal marginal prickles more than 2 mm long; inflorescence mostlv elongate and narrow 5. H. florihunda 1. Hackelia ursina (Greene ex A. Gray) I. M. Johnston Echinospermum ursintiin Greene ex A. Grav, Proc. Anier. Acad. Arts 17: 224. 1882. Lappuki ursina E. L. Greene, Pittonia 2: 182. 1891. Hackelia ur- sina I. M. Johnst. Contr. Gray Herb. 68: 46. 1923. (£. L. Greene, on gravel beds of Bear Can- yon in the Bear Mountains, New Mexico, 4 Octo- ber 1880) Lappula leucantha E. L. Greene, Leafl. Bot. Observ. Grit. 1: 152. 1905. (O. B. Metcalfe 1475, Shady Canyon of Iron Creek, Black Range, Grant County, New Mexico, 11 October 1904) = var. ursina L. pustulata Macbride, Contr. Gray. Herb. 48; 39. 1916. Hackelia ursina var. pustulata J. L. Gentry, Southwestern Naturalist 19(2): 144. 1974. (C. G. Pringle 563, hills west of Chihuahua, Mexico, 23 October 1885) L. heliocarpa Brand, Feddes Repert. Spec. Nov. Reg- ni. Veg. 18: 310. 1922. Hackelia heliocarpa Brand, Pflanzenr. IV 252 (Heft. 97) 120. 1931. (C. G. Pringle 2004, Canyon below Cusihuiriachic, Chihuahua, Mexico, 21 September 1888) = var. pustulata Hackelia ursina var. diaboli J. L. Gentry, South- western Naturalist 19(2): 143. 1974. (G./. Harri- son 1880, Devil's Canyon, Pinal County, .Arizona 16 May 1926) = var. diaboli Stems erect, 1-several, sometimes branched near the base, 3-14 dm tall, hispid or hirsute with spreading bristles or often ap- pressed strigose also, especially above; leaves at the base of plant oblanceolate, long petiol- ate, obtuse, 2.5-14 cm long, 5-15 mm broad, hi.spid-hirsute, pustulate, the cauline leaves gradually reduced above, oblanceolate to narrowly ovate, broader than the basal ones; inflorescence open and spreading; bracts evi- dent throughout the cymes; calyx 1.5-3.5 mm long in fruit, the segments oblong to lan- ceolate, hispid; pedicels 1.5-10 mm long; co- rolla white or tinged with yellow, the tube 1.5-2.5 mm long, the limb 5-11 mm wide; fornices evident, papillate; style 0.8-1.8 mm long, longer than nutlet; nutlets 2-3 mm long, ovate to lanceolate, intermarginal prickles present or lacking, marginal prickles 7-11 on each side, slightly connate at the base or fused for half their length into a cupulate wing, dorsal .surface muricate-hispi- dulous to nearly smooth. Gravelly creek beds, rocky terraces, can- yons, and talus slopes or moist areas, 3,500-8,500 feet elevation, mo.stly in the oak, juniper, or pinus communities. Southern Ari- zona, New Mexico, and northern Mexico. May to August. Our plants of H. ursina, as here described, are the only white-flowered .species and can be .separated into three varieties by the fol- lowing key: Nutlets 2-2.5 mm long, with marginal prickles 1-2 mm long; flowering July and August; Pinal Mountains, Arizona, and western New Mexico to northern Mexi- co 2 Nutlets 2.5-3.5 mm long, with marginal prickles 2-3 mm long; flowering in May; rare in Devil's Canyon, Pinal County, Arizona var. diaboli J. L. Gentry December 1979 HuiGINS: B()»A(;iNACEAE OF THE SoUTHWEST 349 2(1). Corolla limb 5-7.5 mm broad; pedicels rarely more than 2.5 mm at anthesis; southwestern New Mexico var. itrsina Corolla limb 7.5-10 mm broad; pedicels mostly more than 3 mm at anthesis; Pinal Mountains, Arizona, and northern Mexico var. piistidata (.Vlacbr.) [. L. (ientrv 2. Hackelia besseyi (Rydb.) J. L. Gentry Lappiila hcssciji Rydb.' Bull. Torrev Club 31: 636. 1904. //. Icptophi/lhi var. bessei/i Brand, Pflan- zenr. IV 252 (Heft. 97): 127. 1931. //. hcssciji C.entry, Southwestern Naturalist 19(2): 1.39. 1974. (C. E. Besseif, mouth of Chevenne Canvon, (Colo- rado, 25 July 1895) L. grisea Woot. & Standi. Contr. U. S. Natl. Herb. 16: 164. 1913. H. grisea I. M. Johnst. J. .Arnold .\rbor. 16: 194. 1935. (E. O. Wooton, James Can- yon, Sacramento Mountains, New Mexico, 6 Au- gust 1905) Steins erect, solitary, 3-11 dm tall, canes- cent with strigose or villous-hirsute hairs; leaves at base of plant oblanceolate, 2-9 cm long, (7)10-17 mm broad, obtu.se, strigose to hirsute-hispid, pustulate, cauline leaves grad- ually reduced above 2-13 cm long, 5-10(12) mm broad; inflorescence open and spreading; bracts evident only near base of cyme; calyx 1-1.5 mm long in fruit, the segments lance- ovate; pedicels in fruit 3-5 mm long; corolla tube 0.8-0.9 mm long, limb 1.5-2.5 mm broad, blue; fornices evident, papillate; style shorter than nutlet; nutlets 2-2.5 mm long, ovate to ovate-lanceolate, intramarginal prickles lacking, marginal prickles 8-13 on each side, distinct or slightly connate, a long and short prickle alternating, dorsal surface muricate-hispidulous. In the foothills, extending to moderate ele- vations in the mountains, 6,000-9,000 feet, in association with Pinyon-Juniper and Fir-Pine stands. El Paso County, Colorado, south through New Mexico to Trans-Pecos Texas. July to September. The very small corollas, with ascending lobes, easily di.stinguish this plant from all other members of Hackelia in North Ameri- ca. 3. Hackelia hirsuta (Woot. & Standi.) I. M. Johnston Lapptiki hirsuta Woot. & Standi. Contr. U. S. Natl. Herb. 16: 164. 1913. //. hirsuta I. M. Johnst. Contr. Gray Herb. 68: 46. 1923. (G. Heller 3793. 9 miles east of Santa Fe, New Mexico, 2 July 1897) Stems 1 or few, often bluish tinged at the base, erect, or widely branched from the base and throughout, 1-8 dm tall, spreading hispid below, hirsute to strigose above; leaves at base of plant oblanceolate, acute, petiolate, withering early, 2-7 cm long, 5-10 mm broad, villous-strigose to hirsute, ciliate on the petioles, moderately pustulate, cauline leaves oblanceolate to linear-oblong, 3-10 cm long, 5-12 mm wide, ciliate; inflorescence open, widely spreading; bracts 3-10 mm long, evident throughout; calyx 2-3 mm long in frviit, the .segments oblong to lanceolate; pedicels 5-10 mm long in fruit; corolla blue with a white eye, the tube 1.5-^ mm long, limb 4-8 mm broad; fornices evident, papil- late; style 0.6-1.1 mm long, shorter than nut- let; nutlets 2.5-3.5 mm long, ovate-lanceol- ate, intramarginal prickles absent, marginal prickles 4-7 on each .side, slightly connate or distinct at the base, dorsal surface mmicate- hispidulous. On dry, open hillsides or shale roadcuts, in oak canyons or coniferous forests, or rarely moist areas, 6,000-10,000 feet elevation. En- demic to north central New Mexico. A striking and very distinct species due to the spreading branches and the con.spicuously hirsute-hispid stems and leaves. 4. Hackelia pinetorum (Greene ex A. Gray) I. M. Johnston Echinospermum pinetorum Greene ex \. Gray, Proc. Amer. .Acad. .\rts 17: 224. 1882. Lappula pineto- rum 1. M. Johnst. Contr. Gray Herb. 68: 45. 1923. H. florihunda var. pinetorum Brand, Pflan- zenr. IV 252 (Heft. 97): 127. 1931. (£. L. Greene. Pinos Altos Mountains, New Mexico, July and September 1880) = var. pinetorum H. pinetorum var. jonesii J. L. Gentry, Southwestern Naturalist 19(2): 142. 1974 {M. E. Jones. Soldier Canyon, Sierra Madre, Chihuahua, .Mexico, 16 September 1903) = var. jonesii Sterns 1 or few, erect, 3-8 dm tall, grayish hirsute below, becoming strigose above; leaves at base of plant withering early, ellip- tic to oblong or oblanceolate, obtuse, petiol- ate, 3-8.5 cm long, 10-20 mm broad, hirsute to hispidulous, cauline leaves reduced up- ward, 3-12 cm long, 8-25 mm broad; in- florescence open and spreading; bracts lack- 350 Great Basin Naturalist Vol. 39, No. 4 ing or 1-2 at the base; calyx 1.5-2 mm long in fruit, the segments lanceolate to oblong; pedicels 2-5 mm long in fruit; corolla pale blue, tube 1.3-1.6 mm long, the limb 4-7 mm broad; style not exceeding nutlet; nutlets 2-3 mm long, lanceolate to lance-ovate, in- tramarginal prickles small, 1-3 or absent, marginal prickles 4-7 on each side, distinct or slightly connate at the base, less than 2 mm long, dorsal surface muricate hispidulous. Moist, shaded places in Douglas-fir or oak woods or pine woodlands at elevations 6,000-9,000 feet. Coconino County, Arizona, south to southeastern Arizona to southern New Mexico and Trans-Pecos Texas, south into Chihuahua, Sierra Madre, Occidentale, Mexico. June to August. The northern phase of H. pinetorum is the most common and is var. pinetorum. The southern element has been called var. jonesii and enters our flora only in the Organ Moun- tains of southern New Mexico. It is distin- guished from the typical plant by the absence of intramarginal prickles; however, there is some introgression between the two varieties in the Organ Mountains. 5. Hackelia floribunda (Lehm.) I. M. John- ston Echinospermtim floribiindum Lehm. Stirip. Pug. 2; 24. 1830. E. deflexum var. florilmndwn S. Wats. Bot. King Exp. 245. 1871. Lappida floribunda E. L. Greene, Pittonia 2; 182. 1891. H. floribunda I. M. Johnston. Contr. Gray Herb. 68: 46. 192.3. {Drummond, Saskatchewan) Lappula Icptophijlla Rydb. Mem. New York Bot. Gard. 1: 329. 1900. H. Icptophijlla I. M. Johnst. Contr. Gray Herb. 68: 46. 1923. (Several speci- mens cited from Montana and Wyoming) Stems stout, erect, 5-12(14) dm tall, reflex- ed or spreading hirsute or strigose below; leaves at base of plant withering early, oblan- ceolate to elliptic-oblong, 4-20 cm long, 5-20(25) mm broad, petiolate, apex obtuse to acute, hirsutulous-appressed, cauline leaves sessile, gradually reduced upward; in- florescence elongate, rather narrow with strongly ascending, many-flowered branches; bracts lacking or 1-2 at base of cymes; calyx in fmit 2-3(3.5) mm long, the segments ob- long to lance-oblong, hirsute; pedicels 1-3.5 mm long at anthesis, in fruit becoming 4-7(10) mm long; corolla blue or rarely whit- ish, the tube 1-2 mm long, the limb 4-7 mm broad; fornices small, obscurely papillate; style shorter than nutlets; nutlets 3-5 mm long, ovate or ovate-lanceolate, intramarginal prickles lacking or rarely present on a few of the nutlets of the inflorescence, marginal prickles 5-8 on each side, distinct or slightly connate, or sometimes fused for half their length, 1.5-3 mm long, dorsal surface with a faint median ridge, muriculate-hirsutulous. Moist to moderately dry places in the mountains or foothills, or along stream banks, associated with oak, aspen, and evergreen forests 4,000-10,500 feet elevation. British Columbia, Alberta, and Sa.skatchewan, south to Nevada, Arizona, and New Mexico. Dis- junct to Durango, Mexico, less often in Washington, Oregon, and California. July to August. There is some variation within H. flori- bunda, such as the fusion of marginal prickles or not, and the presence or absence of in- tramarginal prickles. These phases in the past have been called H. leptophylla; however, they seem to be wholly arbitrary and not worthy of any taxonomic recognition. ANOMIOPSYLLINAE (SIPHONAPTERA: HYSTRICHOPSYLLIDAE), II. THE GENERA CALLISTOPSYLLUS, CONORHINOPSYLLA, MEGARTHROGLOSSUS, AND STENISTOMERA Vernon J. Tipton'. Harold E. Stark, \ John A. Wildie' .'Abstract.— The subfamily Anoniiopsyllinae in North America consists of five genera: Anomiopsylhis Baker; Cal- listopsyUus Jordan and Rothschild; Conorhinopsi/Ua Stewart; Megarthroglossus Jordan and Rothschild; and Stenisto- merci Rothschild. The revision oi AnoniiopsyUns was the subject of a previous paper, and the remaining four genera are treated herein. The North American genus Jordanopsylla Traub and Tipton and the Palearctic genus Wagnerina loff and .\rgyropulo had been included in the subfamily Anomiopsyllinae by loff and Argvropulo, but these two genera possess characters which are not consistent with the current definition of the subfamily. All species are con- sidered to be "nest fleas," but modifications for a nest habitat are more pronounced in some species than others. Diagnoses, descriptions, and keys are given for genera and most species. In addition, illustrations, host records, and geographical distribution are given for each of the 21 species. The family Hystrichopsyllidae Tiraboschi, 1904, consists of nine subfamilies. In four of the subfamilies (Acedestiinae, Ctenophthal- minae, Dinopsyllinae, and Stenoponiinae) genal combs are present. In three additional subfamilies genal combs are present except in certain genv:ra (Hystrichopsyllinae except Atyphloceras, Neopsyllinae except Catallagia and Delotelis, and Rhadinopsyllinae except WenzeUa and TricliopsyUoides). There is no genal comb in the subfamily Anomiopsyl- linae, and it is vestigial or lacking in Listrop- syllinae. In Anomiopsyllinae the eye is re- duced, vestigial, or absent. The pleural arch is absent, the lateral metanotal area is re- duced or absent, the upper anterior margin of the metepisternum is concave, and there is no striarium on abdominal segment II. There is marked reduction in chaetotaxy in Ano- miopsyllus, not marked in Stenistomera, and intermediate in the other three genera of the subfamily. Anomiopsyllinae in North America consists of five genera: AnomiopsyUus Baker; Collis- topsylhis Jordan and Rothschild; Con- orhinopsylla Stewart; Megarthroglossus Jor- dan and Rothschild; and Stenistomera Rothschild. The genus AnomiopsyUus was re- vised by Barnes, Tipton, and Wildie (1977) in the first of a series of papers on the family Hystrichopsyllidae. The remaining four North American genera are discussed in this paper. When Traub and Tipton (1951) described the genus Jordanopsylla, they indicated that it differed from the other genera of Ano- miopsyllinae as follows: "The metephimere is free; the pleural arch is present; the upper margin of the metepisternum is convex; and the internal marginal tubercle of the mete- pisternum is vestigial." Based on these differ- ences they erected the tribe Jordanopsyllini. loff and Scalon (1954) recognized the mor- phological similarities between Jordanopsylla and Wagnerina loff and Argyropulo and placed the latter with Jordanopsylla in the tribe Jordanopsyllini. The characters given above provide ample reason to question in- clusion of these two genera in the subfamily Anomiopsyllinae. EopsyUa Argyropulo, like Wagnerina, is palearctic in distribution and so will not be considered here. The five North American genera of Ano- miopsyllinae exhibit ecological as well as morphological affinities. Their morphological similarities reflect their shared evolutionary history and habitat. They are considered "nest fleas," which means they are better adapted to conditions in the nest than on the 'Department of Zoology, Brigham Young University, Provo, Utah 84602. 'P.O. Box 636, Montello, Nevada 898.30. 'U.S. Army Environmental Hygene Agency, Regional Division-West, Fitzs.mons AMC, Aurora, Colorado A0045. 351 352 Great Basin Naturalist Vol. 39, No. 4 host. Traub (1972) listed characters which ap- pear to be common to nest fleas, and all five genera possess these characters in varying de- grees. The genus AnomiopsyUiis appears to have better adapted to the nest environment than other genera of the subfamily. The out- standing features of AnotniopsijUus anatomy that appear to be adaptive are "complete evelessness, extreme reduction in number of setae and spines, extensive loss and fusion of structures in the metathorax, and loss of structures in both meso- and metacoxae." In contrast to the highly modified nature of the thorax, the abdominal segments are only moderately specialized, though marked by loss of some structures in the male genitalia" (Barnes et al. 1977). Other nest fleas share most of these morphological characters. If AnomiopsijUus represents the extreme in ad- aptation to nest conditions, Stenistomera may represent a more moderate adaptive position in that it has well-developed setation, includ- ing a comb on the meso tibia. Discussion Species density of fleas is much greater in the western half of the United States than in the eastern half. Hubbard (1947) indicated there were 33 genera and 56 species and sub- species of fleas known from east of the 100th meridian, and 67 genera and 230 species and subspecies were known from west of the 100th meridian. In the intervening years since 1947 several taxa have been described and thus these figures have been adjusted up- ward. For example, Benton (in press) lists 67 species and subspecies of fleas in the United States east of the Mississippi River. Of the 33 genera listed bv Hubbard 19 genera occur only west of the 100th meridian. If the line which divides "east" and "west" is moved east 138 miles to the 98th meridian, then 27 genera are exclusively western. By way of comparison, there are two genera that occur only in the east. Dice (1943) emphasized the great diversity of habitats in the western United States in that he lists 15 biotic prov- inces west of the Mississippi River, but only four east of the Mississippi River. Jellison (1979) has pointed out that diversity of habi- tat is reflected in the large number of species of rabbits, squirrels, and other host animals in the west. Simpson (1964) calls attention to a west- east species gradient for mammals and uses the term front to denote an abrupt decline in species density along a species density gradient. He also points out that mean annual precipitation and topographical relief are the two main factors that contribute to fronts. The 100th meridian corresponds roughly with the 20-inch isohyet, and the high relief topography of the west contrasts sharply with the low relief of the plains states and the relativelv low relief of the eastern United States. In Utah, a state of high relief, there are ap- proximately 115 species and subspecies of fleas in an area of about 85,000 square miles. Hopla (1979) estimates there are 40 species and subspecies of fleas in Kansas, a state of low relief with an area of slightly more than 82,000 square miles. Traub (1979) cautioned that "while mean annual precipitation and topographical relief are of course highly important factors, one must also consider the geologic history of the two areas and the origins and dispersals (his- torv, biogeography, phylogeny and evolution) of the respective host and siphonapteran faunas." The distributional pattern of ectoparasites may be influenced by their life-style. Insect parasites which are parasitic during all stages of their life history and are host specific tend to have distributional patterns similar to their hosts. On the other hand, insect parasites with at least one free-living stage and which are not host specific often have distributional patterns somewhat different from those of their hosts. Fleas in the subfamily Anomiop- syllinae are characterized as "nest fleas." There are 35 recognized species and sub- species in the five North American genera, of which four genera (33 species) occur west of the 98th meridian only and one genus {Con- orhinopsylhi, 2 species) occurs only east of the lOOtii meridian. They have been collect- ed from the nests of Xeotoma species and other nest-building rodents; although a few specimens have been collected from species of Ochotona, Spilogale, and other nonrodent hosts. Doceniber 1979 Tipton et al.: Anomiopsyllinae 353 Key to North American Genera of Anomiopsx llinae (Males) 1. Pronotal comb absent; chaetotaxy greatly reduced; eye absent; caudoventral margin of coxae II and III with acuminate spin- Ammiiopsyllus Pronotal comb present; chaetotaxy variable; eye vestigial or absent; caudoventral margin of coxae II and III without acuminate spur 2 2(1). Sternum VIII greatlv modified, spiniforms on distal apex; clasper elongate (Figs. 18, 19, 22, 23) '. Conorhinopsylla Sternum VIII not modified, without spiniforms on distal portion; clasper not elongate 3 3(2). Eye vestigial; sternum VIII covers more than one-half of distal arm of sternum IX; apex of movable process of clasper on a level with or extends only slightly dorsad of immovable process of clasper Megarthroglossus Eye absent; sternum VIII covers less than one-half of distal arm of sternum IX; apex of movable process of clasper extends well beyond apex (dorsad) of immovable process of clasper 4 4(3). Frons evenly convex; dorsal margin of aedeagal apodeme not extended into long coiled rod; preantennal area and abdominal sterna lacking enFarged bristles; mesotibia without dorsolateral comb CallistopsijUufi Frons subacuminate; dorsal margin of aedeagal apodeme extended into long coiled rod; preantennal area and abdominal sterna possessing slightly or greatly enlarged bristles, mesotibia with dorsalateral comb Stenistomera CaUistopsyllus Jordan and Rothschild Calli.stopsylhi.'i Jordan and Rothschild 1915:46 (Type species CuUistopsijUus tcrinus Rothschild 1905 as Cerato- phiiUus)- Ewing 1929:162; Jellison and Good 1942:26; Ewing and Fox 1943:102.111; Hnbbard 1947:274,281; Holland 1949h:55,96; Traub and Tipton 1951:267-268; Traub and Hoff 1951:1-23; Jellison. Locker, and Bacon 1953:17; Hopkins 19.57:64-87; Parker and Howell 1959:597-604; Hopkins and Rothschild 1962:.354; Jellison and Glesne 1967:28-29; Lewis 1974:147-167. CaUistopsijUus was described by Jordan and Rothschild in 1915 and CeratophyUus te- rinus Rothschild, 1905, was designated as tvpe species. Since the original description, three new species have been described: C. deutenis Jordan, C. campestris Holland, and C. pamterinus Wagner, but the latter was subsequently placed as a junior synonym of tcrinus. Our study has demonstrated that morphologic differences are slight and inter- grade in several geographic localities. It is considered advisable to regard CaUistopsyllus as monotypic, with deutenis and campestris as subspecies of C. terinus. CaUistopsyllus tc- rinus is distributed widely over most of west- ern North America. In recent correspond- ence, Traub (1978) expressed doubt that a single species of a nest flea would have such broad distribution. However, morphological characters used to separate described species are within the range of variation found in one locality, especially in New Mexico. The original description of C. deutenis matches the majoritv of specimens collected in east- ern California and western Nevada. Reliable separation is difficult. Even the validity of subspecies is doubted, because the slight dif- ferences are not consistently correlated with geographic distribution. However, sub- specific status is maintained at this time to preserve currently recognized names and to facilitate study of morphological variation. Specimens from Alberta and Sa.skatchewan have been described as C. campestris. At lo- calities in Montana, Wyoming, and .North Dakota, specimens bearing a resemblance to campestris have been found with typical te- rinus. DiACNosis.— There are two rows of pre- antennal bristles in Callistop.syUus and four rows in Stenistomera. CaUistopsyllus may be separated from Megarthroglossus and Con- orhinopsylla by the absence of an eye and from Anomiopsyllus by the presence of a pronotal comb. One of the most dis- 354 Great Basin Naturalist Vol. 39, No. 4 tinguishing characteristics of CallistopsyUus males is the very prominent movable process which greatly exceeds the length of the fixed process and is very large in relation to the comparatively small body. The movable pro- cess has two spiniform bristles, similar to those of Stenistomera and some Anomiop- sylltis but unlike ConorhinopsyUa and Me- garthroglossus, which lack spiniform bristles. Description.— Head: Interantennal su- ture present. Frontal tubercle present. Fron- tal margin evenly rounded. Eye absent. Head bristles not spiniform as in some Stenisto- mera. Two rows of preantennal (or pre- occular) bristles; anteriormost row usually posterior to cibarial pump. Antenna of male enlarged greatly over that of female. Pedicel of antenna of male extends over part of club. Trabecula centralis absent. Antennal fossae extend to beyond posterior border of head. Anteroventral angle (oral angle) of head with sharp, small labrum. Maxillary lobe as in Me- garthroglossus. Labial palp with 4 segments which extend beyond foretrochanter; apex more or less symmetrical; other members of Anomiopsyllinae with shorter labial palpi ex- cept Megarthroghssus which has 5 segments extending well beyond trochanter, with asymmetrical distal segment. Apex of genal lobe acute. Thorax: Pronotal comb with 15-18 (usual- ly 17) teeth on both sides, each sex. Pleural arch absent. Metasternum and metepimeron fused with metepisternum. Metepisternum lacks bristles. Lateral metanotal area absent. Metepimeron with spherical spiracle. Pro- sternum not distinctive, without bristles, triangular portion with apex directed posteri- orly as in AnomiopsyUiis. Mesosternum and mesepimeron with one long bristle each. Me- tanotum with about 11 large bristles on each side, about 4 tiny bristles between these. Me- sonotum with one row of about 4 bristles on each side plus 3 pseudosetae. Legs: Coxae lack spurs as in Anomiop- syUiis; coxa I with one long apical bristle; mesocoxa with partial longitudinal break; each femur with heavy, long, curved bristles at apex; none elsewhere on lateral surface; with small bristles along dorsal margin; meso- tibia lacking comblike arrangement of setae as in Stenistomera. Metatarsus with 4 lateral. curved setae plus medially displaced pro- ximal pair. Abdomen: Segment II with spherical spi- racle. Each abdominal segment with single row of bristles; tergum I with 3-4, terga II-IV with 5, and terga V-VI with 4 long bristles; abdominal tergum VII with 2-3 shorter bristles; 3 antepygidial bristles in both sexes, middle bristle longest, upper bristle shortest. Ventral sternum II with no bristles, sterna III-VII with 2-5 long bristles (2 in male). Abdominal tergum I with 3-5 (usually 4) apical spinelets on each side; abdominal tergum II with 1-4, III with 1-3 (usually 1), IV with 0-1 (usually 0) apical spinelets on each side. Modified Segments— Male: Movable pro- cess extends dorsad beyond fixed process of clasper; with two blunt spiniform bristles. Caudal margin of sternum VIII rounded, not distinctively shaped; does not ensheathe ster- num IX. Sternum IX shaped like neck and head of goose, beak pointed dorsad, with 10-12 short, thick setae directed caudad. Ae- deagal apodeme extends as far cephalad as apex of manubrium or beyond; blade shaped, anterior and posterior ends curved slightly dorsad; neck constricted; dorsal margin with groove which receives ventral margin of manubriimi. Median dorsal lobe small, dis- tinct, as is crochet and apex of sclerotized in- ner tube. Penis rods long, coiled almost one full turn. Modified Segments— Female: Caudal mar- gin of sternum VII curved as in smooth arc, lacking lobes and sinuses. Anal stylet slender, almost 4 times as long as broad; apical seta long, with 2 tiny setae near apex; 1 dorsal, 1 ventral. Spermatheca with oval bulga; no col- lar. Length of hilla and bulga approximately equal. CallistopsyUus terinus (Fox) Ccwtophiilhis tntiiniis dutrluri X Microtiis niordax sierrae X Neotoma nest X Perognathus sp. X Pcromyscus sp. X X X Perotnyscus boylii X Pcrom ysctis crin ittis X X Pcromyscus ercmicus X Pcromyscus manictilatus X X X Pcromyscus m. artcmisiac X Pcromyscus m. gamheli X Pcromyscus m. osgoodi X X Pcromyscus m. sonoriensis X X Pcromyscus truei X X Spcrmophilus cohunhianus X Tcimiasciurus hudsonicus alholimbntus X _^ =a^r: ^_ --r^^ 1 1^ =t }r± r % X Fig. 1. Callistopsyllu December 1979 Tipton et al.: Anomiopsyllinae 357 Female: Caudal margin of sternum VII faintly concave. Bulga of spermatheca about IV2 times as long as wide, shorter than hilla. Distribution.— Arizona: Coconino and Navajo Counties. British Columbia: Eagle Pass, Dawson Falls, Marble Lake. California: Nevada and Siskiyou counties. Colorado: Larimer and Montezuma coun- ties. Idaho: Bingham, Butte, and Jefferson counties. Montana: Beaverhead, Park, and Figs. 2-7. Males, claspers; 2, CaUistopsijUus lerinus ccimpestris; 3, C. t. deuterus; 4, C. t. terimis Sternum L\; .5, C. t. campestiis; 6, C. t. deuterus., 7, C. t. terinus. 358 Great Basin Naturalist Vol. 39, No. 4 Petroleum counties. Nevada: Douglas, Nye, and Washoe counties. New Mexico: Lincoln, Otero, Sandoval, San Juan, Santa Fe, and Taos counties. Oregon: Deschutes, Harney, and Jefferson counties. South Dakota: Cus- ter County. Utah: Beaver, Box Elder, Dag- gett, Iron, Juab, Kane, Millard, Piute, Rich, San Juan, Tooele, Utah, Uintah, Wasatch, and Washington counties. Wyoming: Big Horn, Green River, Lincoln, Park, Sweet- water, and Weston counties. Material Examined.— Arizona: (Coco- nino County) 3$ , 3 $ , ex Peromyscus truei (TC-267), Tuba City, 23-M969; (Navajo County) 25 , ex Peromyscus truei (B778), 31 km SW Holbrook, 26-IV-1949. Colorado: (Larimer Co.) I $ , ex Peromyscus manicu- latus. Rocky Mountain National Park, T-VIL 78; (Montezuma Co.) 2 5 , ex Peromyscus ma- niculatus. Mesa Verde National Park, 26-V- 1962; (Park Co.) I $ , ex Peromyscus manicu- latus, 3.2 km S Fairplay, ll-VIII-1949. Idaho: (Bingham Co.) 1 ? , ex Peromyscus maniculatus (19Y) National Reactor Test Site (AEC-NRTS), 12- VIII- 1966 (BYU); 1 ? , same host, same location, 23-IX-1966 (BYU); 1 $ , 1 ? , ex rodent nest, AEC-NRTS (TRA), 11-X- 1966 (BYU); 2 (5 , 3 ? , ex Peromyscus manicu- latus, AEC-NTRS (33y), 19-11-1967 (BYU); 1 ? , ex Neotoma nest, AEC-NRTS (33Y), 10- III-1967; 15, 1$, ex Peromyscus manicu- latus, AEC-NRTS (19H), 3-Vil-1967 (BYU); 1$, 2 ? , same host, AEC-NRTS (33Y), 20- VIII-1967 (BYU); (Butte Co.) 1 ? , same host, AEC-NRTS (38Y), 19-n-1967 (BYU); 2 <5 , I ? , same host, AEC-NRTS (36HF), 22-III- 1967 (BYU). Nevada: (Nye Co.) 6 5 , 7 ? , ex Peromyscus maniculatus (334), 28 km NE Pahrump (Timber, Rocky Mt.) 19-XII-1949; (Washoe Co.) 1 ? , ex Peromyscus manicu- latus (44-E-16), 19-IV-1944. Oregon: (Jeffer- son Co.) 2 5 , ex Peromyscus sp., Ames Ranch, Madras, 20-V-1958. New Mexico: (Lincoln Co.) 15, 3 ? , ex Peromyscus truei, 3-IV- 1948; (Otero Co.) 3 5, 1 ? , ex Peromyscus truei (D443) 10 km E La Luz, Lincoln Na- tional Forest, 1300 m, 22-III-1949; 35,1?, II km NE La Luz, 1500 m, 24-111-1949; (San Juan Co.) 25, 1$, ex Peromyscus manicu- latus (B778), 73 km E Bloomfield, 1900 m, 18-V-1949; (Santa Fe Co.) 2 5 , ex Per- omyscus maniculatus, (4697) (697) Santa Fe, 2150 m, 2-IV-1953; (Taos Co.) 3 5 , 1 ? , ex Peromyscus sp., Pecos, 22-11-1941, McMurry (Jellison). (Some male specimens from New Mexico and the male from South Dakota are terinus near campestris.) South Dakota: (Custer Co.) 1 5 , ex Peromyscus maniculatus (B1511), 45 km W Custer (1660 m, rock led- ges, pine, juniper) 6-VI-1950. Utah: (Beaver Co.) 1 5 , ex Peromyscus maniculatus. Puffer Lake, 25-VH957, D. M. Allied (BYU); (Box Elder Co.) 1 ? , same host, Yost, George Creek, 9-VII-1957, D. M. Allred (BYU); (Daggett Co.) 2 5, 1 ? , same host, Linwood, 15-VII-1954, C. L. Hayward (BYU); 1 ? , ex Dipodomys ordii, Bridgeport, l-VII-1954, C. L. Hayward (BYU); (Iron Co.) 1 5 , ex Per- omyscus maniculatus. Cedar Break National Monument, 21-Vn-1953, D. E. Beck (BYU); (Juab Co.) 1 5 , same host, Callao, 12-VIII- 1953, Beck, Coffey, Killpack (BYU); (Kane Co.) 55 , 5? , ex Peromyscus truei (182), NAV-KAI, 2-XII-1971 (BYU); 2 ? , ex Per- omyscus crinitus (228), same location, 11-XII- 1971 (BYU); (Millard Co.) 2 5 , 1 ? , ex Per- omyscus maniculatus, Fillmore, ll-VII-1952, Killpack and Coffev (BYU); (Piute Co.) 1 ? , same host, Marysvale, 27-VI-1952, Killpack and Beck (BYU); (Rich Co.) 1 ? , same host, Monte Cristo R.S., 24-VI-1953, D. E. Beck; (San Juan Co.) 1 5 , ex Peromyscus sp., VII- 1946; (Tooele Co.) 1 5 , ex Peromyscus ma- niculatus, Dugway Proving Ground (39), 2- Vm-1951; (Utah Co.) 1 5 , same host, Goshen Springs, lO-IX-1965, W. J. Despain (BYU); (Uintah Co.) 15, 1 ? , same host, Jensen, 9- VM953, D. E. Beck (BYU); (Wasatch Co.) 2? , same host, Soapstone R.S.. 31-VII-1948, Mulaik (BYU); 1 ? , same host, Wallsburg, 2- VII- 1953, D. E. Beck (BYU) ex Peromyscus maniculatus osgoodi, 15, 2 $ , Strawberry Valley, ll-VI-64, K. B. Cox (BYU); (Washing- ton Co.) 2 ? , ex Peromyscus eremicus, Graft- on, 17-XII-1950. Wyoming: (Big Horn Co.) 1 ? , ex Peromyscus nuiniculatus (B-2762), 13 km NW Greybull, 7-VI-1940; (Green River Co.) 1 5 , 1 ? , ex Peromyscus maniculatus (G- 862), 31.6 km NW Green River, 1908 m, 28- VI- 1949; (Lincoln Co.) 1 ? , ex Peromyscus maniculatus, Cumberland, 23-VII-1955; (Park Co.) 15, 1 ? , ex Peromyscus manicu- latus (C-2846-5), Yellowstone National Park, Old Faithful Lodge, 22-V-1940; 15 (Finger resembles both terinus and campestris.), ex December 1979 Tipton et al.: Anomiopsyi.linae 359 Penmiy.sciis t)unucuhitus (G-998), 41.7 km NW Cody, Shoshone National Forest, 2431 m (pine, cedar), ll-VllI-1949; (Sweetwater Co.) 1 ? , ex Peromyscus boijlii (B-1642), 17-VIII- 1938; (Weston Co.) 1 S (Finger resembles both terinus and campestris.), ex Peromyscus manicuUittis (F-848), 10 km N Newcastle, 1385 m (pine bhiffs), 26-V-1949. Host synonymy.— Tamiasciurus douglasi albolimhatus = Tamiasciunis hudsonicus al- hoUmhatus. CaUistopsyUus terinus deuterus Jordan, new combination Figs. 1,3,6.8, 11, 14,36,97 CoUistopst/lhis deuterus Jordan 1937:266; Hubbard 1940:37(4); Augiiston 1942a:140; Auguston, 1942b: 150; Jellison and Good, 1942:26; Ewing and Fox, 1943:111; "Hubbard 1943:1-12; Hubbard 1947:283; Hubbard 1949:126; Jellison, Locker, and Bacon 1953:17; Hopkins and Rothschild 1962:358; Beck 1966:76; Beck and Allred 1966:13; Stark and Kinnev 1969:287-294; Lewis 1974: 147-167; Nelson and Smith 1976:51-61. Type host.— Peromyscus sp. Type locality.— Big Bear Lake, Califor- nia. Type specimens.— United States National Museum, Washington, D.C. Diagnosis.— Male: Finger triangular; broad toward base with a narrow, round apex. Anterior margin convex with a rounded angle just above base. In terinus and camp- estris this angle is at base and is sharper. Caudal margin of finger of deuterus some- times convex in apical half. Caudal margin of distal arm of sternum IX with rounded angle about midway, making apex appear narrower than in terinus or campestris. Small notches, thick bristles, and sharper angles found mid- way along the caudal border of distal arm of sternum IX of few specimens of terinus are not present in deuterus specimens available for examination. Apex of distal arm narrower and longer than either terinus or campestiis, especially campestris. Female: Characters are not sufficiently distinctive to separate deuterus from camp- estris or terinus. Distribution.— Arizona: Yavapai Coun- ty. California: Fresno, Mariposa, Mono, Sis- kiyou, and Tuolumne counties. Colorado: Montezuma County. Mexico: Baja Califor- nia. Nevada: Nye Comity. Orecon: Jefferson County. Material examined.— Arizona: (Yavapai Co.) 1$ , e\ Peromyscus truei (B-708), 16.7 km N Prescott, 5-IV-1949 (The male is defi- nitely deuterus, but other Arizona males are terinus (q.v.). The females from Yavapai Co. may be either terinus or deuterus.); 1 $ , ex Peromyscus truei (B-717), 8.3 km N Prescott, 1662 m, 7-IV-1949; 1 ? , ex Peromyscus sp. (A-301), 22-IV-1938. Baja California: 1 $ , 1 ? , ex Peromyscus truei, 1.6 km S. El Con- dor Hwy #2, 31X11-1962, W. J. Wrenn (Traub Collection). California: (Fre.sno Co.) 1 ? , ex Eutamias speciosus (16097), Hunting- ton Lake, 9-VI-1979; 1 $ , ex Peromyscus ma- niculatus (16099), Huntington Lake (Bureau Vector Control, CA); (Mariposa Co.) 1 ,5 , ex Peromyscus sp.. Jet. Yo.semite Creek Trail & Tioga Rd., Yo.semite National Park, 2215 m, 2- VI- 1959; (Mono Co.) 1 <5 , ex Peromyscus maniculatus, Tioga Pass Resort, Jet. Tioga Rd. & Saddle Bag Lake Rd., Invo National Forest, 3077 m, 8-IX-1961; (Tuolumne Co.) 1 (5 , ex Peromyscus maniculatus, Tuolumne Meadows, Yosemite National Park, 2615 m, (Plague Lab); County not given: 1 $ (16478), locality not given (Bureau Vector Control, CA). Nevada: (Nye Co.) 1 $ , ex Peromyscus crinitus. Mercury (AEC-NRTS), 9-XII-1961 (BYU). Oregon: (Jefferson Co.) (See C. t. ma- terial examined). CaUistopsyUus terinus campestris Holland, new combination Figs. 2, 5. 9, 12, 14, 98 CaUistopsifUus campestris Holland, 1949b:98; Hopkins and Rothschild, 1962:358; Senger, 1966:106; Jelhson and Senger, 1973:19; Lewis, 1974:147-167. Type host.— Peromyscus maniculatus os- goodi. Type locality.— Medicine Hat, Alberta, Canada. Type specimens.— Canadian National Col- lection, Ottawa, Canada. Diagnosis.— Male: Finger somewhat rec- tangular or oval, depending upon specimen. Narrower basally and broader distally than in either terinus or deuterus; sides subparallel. Angle of anterior border at ba.se variable; its outline is between that of terinus, which is acute and at the base, and deuterus, which is 360 Great Basin Naturalist Vol. 39, No. 4 rounded and above the base. Distal arm of sternum IX gradually curved along curved margin; apex broadly rounded— more so than in terinus or deutenis. Bristles along caudal border of distal arm of sternum IX about as numerous (13-14) as terinus. Female: Characters are not sufficiently distinctive to separate terinus from deuterus or carnpestris. Distribution.— Alberta, Saskatchew^an. Montana: Big Horn, Custer, and Park coun- ties. Wyoming: Teton County. Material examined.— Alberta: I $ , 1 $ , ex Peromyscus maniculatus osgoodi. Medi- cine Hat, 6-VI-40 (Holland). Saskatchewan: IS ex P. m. osgoodi, Estevan, 28-VI-42 (Hol- land). Montana: (Custer Co.), 1 ? , ex Dipo- domys ordii (C-2574), 45 km S of Miles City, 22-III-1940; 1 5 , ex Peromyscus maniculatus (C-2581-5), 28-III-1940; 1 ? , ex Peromyscus maniculatus (C-2589-S), 29-III-1940; 2 <5 , ex Peromyscus maniculatus (C-2599), 42 km SW of Miles City, 30-III-1940; (Park City) 2$, 1 ? , ex Peromyscus sp., (D-667), 29-VI-1938; (Prairie Co.) \ $ , 1 ? , ex Peromyscus ma- niculatus (E-760), 8 km E Terry (sage 677 m) 26-V-1949. Conorhinopsylla Stewart ConorhinopsyUa Stewart 1930:178; Fox 1940:29, 41; Jellison and Good 1942:42; Ewing and Fox 1943:73; Jelli- son 1945:109-111; Hubbard 1947:274,293; Holland 1949b:55, 101; Traub and Tipton 1951:267-268; Jellison, Locker, and Bacon 1953:37; Hopkins 1957:64-87; Stark 1959:97; Hopkins and Rothschild 1962:359; Jellison and Glesne 1967:67; Lewis 1974:147-167. Diagnosis.— ConorhinopsyUa is the only genus in the subfamily in which the antero- ventral angle of the head is acuminate, there is no clypeal tubercle, there are two rows of bristles on the terga, and the pleural arch is vestigial. The eye is vestigial as in Megarth- roglossus, but in the latter genus there are no spiniform bristles on the genitalia. Male ster- num VIII is greatly modified and bears spin- iform bristles on the distal portion. Description.— Head: Anterior margin of head slightly convex, frontal tubercle absent, oral angle a nipplelike protuberance. Genal process with undulating anterior margin, apex subtruncate to truncate. Two prominent ocular bristles near anterior margin of genal process. Irregular row of 5 preoccular bristles. Eye vestigial, triangular, lightly pig- mented. Trabeculum centralis (area commu- nis) absent. Antennal fossae do not extend to vertex to form interantennal suture; anterior margin with 3-4 small bristles; posterior mar- gin with 4-5 small submarginal bristles. First and second antennal segments with several small bristles plus 5-6 small bristles. Occipit- al area with two large bristles; placoids on both frontal and occipital areas. Maxillary palp 4-segmented. Labial palp 5-8 seg- mented, extends beyond foretrochanter, apex asymmetrical. Genal comb absent. Thorax: Pronotal comb with 12 very broad teeth. Pleural arch absent. Pleural rod very thin and elongate. Legs: Sexual dimorphism in chaetotaxy of hind legs pronounced in S. stanfordi, less so 9 L__.^^— -"^^ 10 Figs. 8-10. Males, sternnni VIM; 8, Callistopsiilliis terinus dculinis: 9. ('. i. raiiipcstris: 10. C. t. terinus December 1979 Tipton et al.: Anomiopsyllinae 361 Figs. 11-13. Males, aedeagiis: 11, CaUistopsijUns tcriniis deuterus; 12, C. /. campcstris; 13, C. t. terinus. 362 Great Basin Naturalist Vol. 39, No. 4 b DISTRIBUTION OF THE GENUS CALLISTOPSYLLUS ■ deuterus • termus campestris Fig. 14. Distiilnition of Callistopsijllns tcriniis .s.sp. December 1979 Tipton et al.: Anomiopsyllinae 363 in S. nidicola. Four pairs of lateral plantar bristles and proximal median pair of bristles on fifth tarsal segments of all legs. Abdomen: Abdominal tergal spinelets vari- able, generally terga I-III with 2 spinelets per side, tergum IV with one per side, terga V-VII without spinelets; each tergum with 1 row of bristles usually consisting of 5 to 7 long bristles intermixed with several minute bristles, ventrad most long bristle shifted an- terad on main row of bristles. Lateral bristles on abdominal sterna variable, usually in row of 2-4 per side. Number of antepygidial bristles 2 or 3 depending on species: with 3, centermost longest, remaining 2 vary from minute to small in males or from two-thirds to three-fourths length of middle bristle in fe- males; with 2, dorsal bristle longest, ventral bristle varies from minute to medium in males, from three-fourths to sube(jual in length to dorsal bristle in females. Modified segments— Male: Fixed process of clasper greatly enlarged, modified in C. stan- fordi, less so in C. nidicola. Sternum VIII re- duced, with apical or subapical spiniform bristles. Sternum IX reduced, with apical and subapical bristles. Details of aedeagal mor- phology vary according to species. Modified segments— Female: Caudal mar- gin of sternum VII convex. Bulga of sperma- theca somewhat beanlike, hilla lacking stria- tions, otherwise lacking sharp line of demarcation between bulga and hilla. Key to The Species of Conorhinopsylla Males Apical portion of distal arm of sternum VIII with 8 or more spiniform bristles (Fig. 19); movable process of clasper extends far beyond apex of immovable process of clasper (Fig. 23) nidicola Apical portion of distal arm of sternum VIII with 5 or fewer spiniform bristles (Fig. 18); movable process of clasper does not extend far beyond immovable process of clasper (Fig. 22) stanfordi Table 3. Distribution of species of Conorhinopstjlhi. Table 4. Host associations of the genus Con- orJtinopsyUa. Host Species Host Species Illinois Indiana Iowa Kansas Maryland Michigan New York Ohio Pennsylvania Utah ' Wisconsin Ontario Glaticomijs sp. Glaiicomijs sabrinus Glaucomys sabrinus macrotis Glaucomijs vohins Gkiucomys voliins colons Neotoma sp. Neotoina floridana Neotoma floridana osagensis Peromyscus sp. Peromyscus maniculatus Procyon lotor Sciuriis carolinensis Sciurus niger Spennophihis townsendi Tamiasciurus hitdsonicus Tamiasciurus hudsonicus loquex 364 Great Basin Naturalist Vol. 39, No. 4 Fig. 15. ConoiIiinopsijUa stanforcii; Male, head and thorax inchiding co.xae. ConorhinopsyUa stanfordi Stewart Figs. 15-18, 20, 22, 24, 26, 31 ConorhinopsyUa stanfordi Stewart 1930:178-179; Jor- dan 1933:267; Fox 1940:41; Hubbard 1940:37(4); Fuller 1942:137; Jellison and Good 1942:42; Ewing and Fox 1943:74; Fuller 1943:5; Jameson 1943:177; Stanford 1944:174; Jellison 1945:109; Hubbard 1947:102; Jellison, Locker, and Bacon 1953:37; Benton 1955:1.39-140; Bur- butis 1956:782; Layne 1958:162; Geary 1959:355; Stark 1959:97; Benton and Cerwonka 1960:383-391; Hopkins and Rothschild 1962:.360; Benton and Smiley 1963:4; Os- good 1964:29-33; Jellison and Glesne 1967:65-66; Bent- on 1967:150-160; Humphreys 1967:188; Whitaker and Corthum 1967:4.32; Holland and Benton 1968:2,56; Tip- ton and Saunders 1971:18; Amin 1973:110-111; Jenkins and Grundmann 1973:76-86; Haas and Wilson 1973: 302-314; Lewis 1974:147-167; Jack.son and Defoliart 1976:.351-356. Type host.— Tamiasciurus hudsonicus. Type locality.— Ithaca, Tompkins Co., New York. Type specimens.— Collection of M. A. Stewart. Diagnosis.— ConorhinopsyUa stanfordi may be distinguished from C. nidicola, the only other species in the genus, on the basis of the following set of characters; eye vesti- gial but sufficiently pigmented to be readily discernible; labial palp with 5-6 segments; in the male the metatibia and first two metatar- sal segments with long prominent bristles; clasper projected caudally, fixed and movable processes of clasper about same size; distal arm of sternum VIII narrow but subapical area slightly enlarged and with 4-5 spiniform bristles; the distal arm of sternum IX is nar- row, tapers apically and bears several long subapical bristles. Distribution.— Illinois: Madison Coun- ty. Indiana: Vigo County. Iowa: Dubuque County. Maryland: Frederick County. Michigan: Clinton County. New York: Otesgo and Ulster counties. Ohio: Athens and Jackson counties. Pennsylvania: Brad- ford and Erie counties. Utah: Sevier County. Wisconsin: Adams and Iowa counties. Canada: Ontario. Material examined.— Maryland: (Fred- erick Co.) I S ex Tamiasciurus hudsonicus loquex (nest), 1.5 miles N Wolf.sville, 28-XI to 3-XII-1970, Traub and Schlitter; 1 ? ex Glauconiiis v. volans, same collection data; December 1979 Tipton et al.: Anomiopsyllinae 365 Figs. 16-17. ConorhinopsyUa stanfordi. me tathoracic legs; 16, female; 17. male. 366 Great Basin Naturalist Vol. 39, No. 4 fo^^2^'niJ:^:!:' ^'"""" ''"^^ ''- ConorJ,nopsy„a stanfor,, ,9. C nidicola. 20-31. ster.n.m IX; 20, C. ./,/„- December 1979 Tipton et al.: Anomiopsyllinae 367 1 $ ex Sciunts carolinensis, Oxon Hill, 5-XII- 1923, E. Chapin. Ohio: (Athens Co.) 1 $ ex nest of flying squirrel, Waterloo Twp., 10-X- 1964, H.'G.'Huinphreys. New York: (Chau- tauqua Co.) 1 (5 ex nest of Glaucomys volans, 1 mile S Fredonia; (Otesga Co.) 1 (5 , 1 ? ex Glaucomys volans, Middlefield Twp., 6-XII- 1956, P. Connor. Discussion.— Stanford (1944) reported C. stanfordi from Utah but this record is very likely in error. If so, then it can be said that C. stanfordi occurs on arboreal squirrels east of the Mississippi River. ConorhinopsyUa nidicola (Jellison) Figs. 19, 21, 23, 25, 27, 31 ConorhinopsyUa nidicola Jellison 1945:109; Holland 1949b: 102; Jellison, Locker, and Bacon 1953:37; Rainey 1956:535-646; Poorbaugh and Gier 1961:201; Hopkins and Rothschild 1962:362; Jellison and Glesne 1967:65; Lewis 1974:147-167. Diagnosis.— ConorhinopsyUa nidicola may be characterized as follows: eye vestigial and not readily discernible; labial palp with 8 or more segments; in the male there is no comb- like row of bristles on the hind tibia, and the tarsal bristles are not so long as in C. stan- 23 Figs. 22-23. Males, claspers; 22, Conorliinopsylla stanfordi; 23, C. nidicola. 368 Great Basin Naturalist Vol. 39, No. 4 Figs. 24-25. Males, aedeagus; 24, Conorltiuopsylhi stdnfordi- 2.5, C. nidocohi. December 1979 Tipton et al.: ANOMiopsvLLir 369 fonli; there is no caudal projection of the male clasper and the movable process of the clasper extends dorsally far beyond the fixed process of the clasper; the distal arm of ster- num VIII is broad and with 8-10 apical and subapical bristles; the distal arm of sternmn IX is broad and with a truncate apex and sev- eral marginal and submarginal bristles. Female characters are not sufficiently dis- tinctive to be diagnostic. Type host.— Neotoma floridana. Type locality.— Lawrence, Douglas Co., Kansas. Type specimens.— Snow Collection, Uni- versity of Kansas and U.S. National Museum. 28 30 Figs. 26-30. Females, sternum VIII; 26, Conorhinopsylla stanfordi: 27. C. nidicola; 28, Stenistonwra hubbardi: 29. S. macrodactyla; 30, S. alpina. 370 Great Basin Naturalist Vol. 39, No. 4 Distribution.- Kansas: Douglas, Riley, and Republic counties. Material examined.— Kansas: (Douglas Co.), 3 (? , 3 ? (paratypes) ex nest of Neotoma sp., Lawrence, ll-XI-1944, R. H. Reamer; 2$ ex Neotoma sp., 1959, A. El-Wailly; 1 ? ex Neotoma floridana, 1959, A. El-Wailly; 1 $ ex nest of Neotoma sp., Lawrence, 10-V- 1948, C. E. Hopla; 1 ? ex nest of Neotoma floridana osagenis, Lawrence, 13-XII-1947, NORTH AMERICA DISTRIBUTION OF THL GENUS CONORHINOPSYLLA 'C. nidicola ■C. stanfordi ^-=^=q>^ Fig r. 3L Distribution oi Cotwrhiuojmjlla nkUcohi and C. slanfordi. December 1979 Tipton et al.: Anomiopsyllinae 371 C. E. Hopla; 1 (5 ex Neotoma floridana, Law- rence, 3-X11-1949, C. E. Hopla; 1 (5 , 1 $ ex nest of Neotoma sp., Lawrence, 30-1X-1948, C. E. Hopla; 1 ? ex nest of Neotoma florid- ami osagensis, 22-X1-1952, D. A. Crossley. Discussion.— ComirhmopsijUa nidicola is associated with Neotoma floridami west of the Mississippi River in Kansas. MegartJiroglossus Jordan & Rothschild Megarthroglos.sus Jordan and Rothschild 1915:46 (Type species: Mcgarthroglos.'ius procu.s Jordan and Rothschild, 1915); Ewing 1929:162; Jellison and Good, 1942:83; Ewing and Fox^ 1943:113; Hubbard 1947:296; Holland 1949:89; Eads 1950:53-54; Traub and Tipton 1951:267-278; Jellison, Locker, and Bacon 1953:104; Traub 1953:77-85; Morlan 1954:446; Mendez 1956: 159-192; Hopkins 1957:64-67; Finlev 1958:213-552; Parker and Howell 1959:597-604; Stark 1959:98; Hop- kins and Rothschild 1962:365; Holland 1965:1052; Jelli- son and Glesne 1967:16; Stark and Kinnev 1969:287-294; Mendez and Haas 1972:285-288; Lewis 1974:155; Nel- son and Smith 1976:54; Egoscue 1976:475-480. The Nearctic genus Megarthroglossus was established by Jordan and Rothschild in 1915 with Megatihroglossus procu.s as the type spe- cies. Baker (1895) described Pulex long- ispinus, which was subsequently placed in the genus Ceratophijllm by Wagner (1898) and in the genus Megarthroglossus by Jordan and Rothschild (1915). Mendez (1956) revised Megarthroglossus and included 12 species and 1 subspecies. Since this revision an additional 3 species have been described: M. cav- ernicohis Mendez and Haas, 1972; M. wilsoni Mendez and Haas, 1973; and M. weaveri Eads and Campos, 1977. Megarthroglossus muiri was reduced to a subspecies of M. pro- cus by Hopkins and Rothschild (1962), and we have cho.sen to retain the two subspecies as valid taxa even though they are very sim- ilar morphologically. In this paper M. pyg- maeus (Wagner 1936) is considered a junior synonym of M. s-penceri, and M. divisus exse- catus is reduced to a junior synonym of M. divisus. As a result of these taxonomic changes, the genus Megarthroglossus includes 13 known taxa, of which one is a .subspecies. The genus Megarthroglossus has been col- lected in a variety of habitats and from manv hosts (Table 6). Species of the genus Neototna appear to be preferred hosts. Species of the genera Eutamias, Glaucomys, Ochotona, Per- omyscus, and Tamiasciurus are considered to be secondary hosts. The geographical distribution of species and subspecies of Megarthroglossus is given in Table 5 and Figures 100, 101. Specimens have been collected from Texas to Canada and from California east to Nebraska. Diagnosis.— Megarthroglossus may be sep- arated from Callistopsyllus and Anomiop- Table 5. Distribution of species and subspecies of Megarthroglossus n »-. Taxa U U M. hecki M. bisetis M. cavcrnicohis M. divisus M. jaiucsoni M. procus Diuiri M. procus procus M. sicctnius M. sierrae -M. smiti M. spenceri M. weaveri M. wilsoni \ \ \ X X X X X X X X X X X X X XX XX X XXX X X X X X X 372 Great Basin Naturalist Vol. 39, No. 4 syllus on the basis of long labial palpi which extend beyond the procoxa and trochanter. Conorhinopsylla and Stenistomera, the re- maining North American genera of the sub- family Anomiopsyllinae, like Megarthro- glossus, possess long labial palpi, but, unlike Megarthroglossus, they have spiniforms on the male genitalia. Description.— Head: Anterior margin of head evenly convex in females; more elo- ngate in males, anterior margin convex to subangulate. Frontal tubercle (clypeus) pres- ent, small, inconspicuous. Eye present, reduc- ed, lightly pigmented. Maxillary lobe acumi- nate to subacuminate, normally not extending beyond middle of second segment of labial palp. Maxillary palp 4-segmented. Labial palp 5-segmented and extends one- half to one full segment beyond procoxa. Tra- beculum centralis absent. Genal lobe evenly convex. Genal comb absent. Occvilar bristle in row of 4 alternating long and short, pro- ceeding from eye to genal margin. Pre- occular bristles 0-4, (usually in row of 3), di- rectly over cibarial pump. Posterior antennal fossae with 2 long plus several minute bristles. Antennal fossae prolonged in males, may or may not extend to vertex, to form Table 6. Host associations of the genus Megarthroglossus. Hosts Ammosperrnophihts leucurus leuciirus Canis latrans "Chipmunk" Dipodatnys spectabilis Eutamias sp. Eutamias minimus Eutamias quadrivittatus Eutamias totvnsendi Gkiucomys sp. and nest Glaucomijs sabrinus Glaucomys s. lascivus Glaucomys s. oregonesis Lynx sp. Microtus longicaudus "Mouse" Neotoma sp. and nest Neotoma albigula and nest Neotoma cinerea and nest Neotoma c. acraia Neotoma c. cinerea Neotoma c. occidentalis Neotoma lepida Neotoma I. lepida Neotoma mcxicana Neotoma micropus Neotoma m. canescens Onychomys leucogaster December 1979 Tipton et al.: Anomiopsyllinae 373 falx. Tentorial arch present, well developed, located anterior to eye. Thorax: Prosternuni with dorsal angnlate to siibangulate inflexion at posterior one- third of ventral margin. Pronotal comb pres- ent, 6 to 8 blunt, broad spines per side. Tho- racic link plates II and III well developed. Mesonotum with 3 distinct rows of bristles. Mesepimeron usually with 3 long bristles, an- teriormost close to or directly over pleural rod. Mesepisternum with row of small bristles, extending from dorsal angle to mid- segment. Metanotinn with one row bristles. Lateral metanotal area not complete, scle- rites partially fiLsed. Metepisternum partly fused with metasternuni and nietepimeron to metanotum; setation sparse, normally with 3 long bristles, one near caudal margin, 2 in proximity to pleural rod. Legs: Profemur with 2-3 lateral bristles on mesal surface. Metatarsus II with 1-2 long, lateral, apical bristle(s), extending from one- half to full length of metatarsus V. Meta- tarsus V with 4 pairs lateral plantar bristles, 1 basoventral pair. Abdomen: Abdominal tergal spinelets vari- able, generally tergum I with 1-2, tergum II with one per side. Abdominal terga with one Table 6 continued. Ochotoiui sp. Ochotona princeps Oclwtona schi.'iticcps Ochotona miiiri Pewmijscus sp. Pcwmijsctts hoijlii Perom ijscits crin itus pergmciUs Pewmijscus leucopus Pewmijsciis maniciihitus Pcromijsciis in. nihidus Peiontyscns in. .sonoricnsis PeroinyscHs nasitttis Peromyscus trtiei Pcroniy.scus t. nevadcnsis Rattits norvcgicus Sigmodon hispidus Sorex pcdiistiis navigator SpHogalc sp. Spilogah' gracilis ■saxatilis Syhilagiis nuttallii Taniiasciiiriis sp. and nest Tamidscitirus hudsoniciis Taniia.^ciiinis /i. douglasi Taininsciurus h. alholinibatus Tamiascitirus It. freinonti Tamiascitirus h. strcatori Thomomys bottae TJiomomys talpoides "Weasle" X X X 374 Great Basin Naturalist Vol. 39, No. 4 row of 5-7 long bristles intermixed with sev- eral minute bristles, ventralmost long bristle shifted anterad of main row of bristle. Lateral bristles on abdominal sterna variable, usually in row of 2-4 per side. Antepygidial bristles 2-3 depending on species: those with 3, cen- termost longest with upper and lower varying from minute to small (in males) or from two-thirds to three-fourths length of center bristle (in females); those with 2, up- permost longest with lower varying from minute to medium (in males), three-fourths to almost as long as upper (in females). Modified segments— Male: Fixed process of clasper with hump (inner fovea), one-half dis- tance from acetabular area to apex of pro- cess, may be marginal or submarginal. Caud- al margin of fixed process with 2-3 (usually 3) long, stout bristles dorsad of acetabular area, one bristle ventrad to acetabular area. Shape of caudal margin of stenuim VIII vari- able, diagnostic in some species, normally en- sheathing distal one-half to two-thirds of dis- tal arm of sternum IX. Proximal arm of sternum IX expanded, with or without ceph- alad-directed projection. Shape of distal arm of sternum IX variable, diagnostically distinct in some species. Aedeagal apodeme blade shaped, tapering at neck, without proximal or apical spur. Dorsal margin either straight, with slight undulation, or possessing promi- nent hump in proximity of crescent sclerite. Median dorsal lobe acuminate, truncate, or broadly convex. Crochet of aedeagus large, conspicuous; caudal border variable; apical process hooklike, rounded, or absent. Apex of sclerotized inner tube truncate, with distinct dorsolateral projection. Satellite sclerite closely associated with sclerotize inner tube. Penis rods uncoiled, extending slightly dorsad of apex of aedeagal apodeme. Modified segments— Female: Caudal mar- gin of sternum VII undulate, sinus present or absent on ventrolateral aspect. Anal stylet variable in length, setation varies according to species. Spermatheca variable, bulga may be elongate or compressed. Hilla bent sharp- ly or gently curved dorsad, distal end varying from narrowly convex or broadly convex to subtruncate. The genus Megarthroglossus is a complex group of closely related species with many similar morphological features. Often the de- scription of one species applies equally well to several other species. For this reason, a de- tailed description of all species is not given but, instead, a short, narrative diagnosis for each species is presented. Keys to both males and females (except for tlie female of M. sicamus) are given. It should be emphasized that overlap between species was observed for several taxonomic charac- ters. Characters used in the key include mea- surements whenever possible with the intent of excluding such amlDiguous terms as promi- nent, not deeply indented, and narrow, to mention but a few. In addition, the key to the females may not be entirely satisfactorv, but it is intended to fill a void because current literature does not include keys to the fe- males. Megarthroglossus sicamus is not in- cluded in the key because specimens were not available. If M. sicamus were included in the key, it would be placed near M. jamesoni and M. cavernicolus. Key to Species of Megarthroglossus (Males) 1. Height of hump on dorsal margin of aedeagus greater than 10 microns 2 Hump of dorsal margin of aedeagus absent, or, if present, height less than 10 microns 6 2(1). Posterior margin of sternum VIII evenly convex; inner fovea of inunovable process more than 60 microns below dorsal margin; hump on dorsal margin of aedeagus exceeds 30 microns sicamus Posterior margin of sternum VIII sinuate; inner fovae of immovable process less than 60 microns below dorsal margin (Fig. 32); hump on dorsal margin of aedeagus variable 3 December 1979 Tipton et al.: Anomiopsyllinae 375 Anterior margin of finger of clasper with angular denticle; hump on dorsal margin of aedeagus exceeds 20 microns; inner fovea of immovable process ex- ceed 50 microns below dorsal margin; ventrolateral lobe of sternum VIII evenly convex spctueri Anterior margin of finger of clasper without angular denticle; inner fovea of immovable process less than 50 microns below dorsal margin; ventrolateral lobe of sternum VIII variable 4 4(3). Median dorsal lobe of aedeagus reduced to short, blunt lobe; posterior margin of sternum VIII sinuate, with tnmcate to subtruncate ventrolateral lobe Median dorsal lobe of aedeagus long, with curved apex; posterior margin of sternum VIII undulate, with a short subacuminate lobe sniiti 5(4). Hump on dorsal margin of aedeagus less than 20 microns; length of labial palp segment V less than 220 microns; inner fovea of immovable process exceeds .30 microns, marginal; ventrolateral lobe of sternum VIII subtrrmcate sierrae - Hump on dorsal margin of aedeagus exceeds .30 microns; length of labial palp segment V exceeds 220 microns; inner fovea of immovable process exceeds 30 microns, submarginal; ventrolateral lobe of sternum VIII truncate ,.. jainesoni 6(1). Ventrolateral lobe of sternum VIII long, fingerlike, curved ventrad; inner fovea of immovable process exceeds 50 microns, marginal becki Ventrolateral lobe of sternum VIII not fingerlike; posterior margin of sternum VIII variable; inner fovea of immovable process less than 50 microns, submarginal 7 7(6). Posterior margin of sternum VIII evenly convex; 2 or 3 antepvgidial bristles 8 Posterior margin of sternum VIII sinuate, with variable shaped ventrolateral lobe; 3 antepvgidial bristles 8(7). Two or 3 antepygidial bristles per side; inner fovea of immovable process less than 35 microns; segment V of labial palp less than 220 microns in length; cro- chet spur exceeds 10 microns in length; finger of clasper less than 120 microns in length; metatarsal segment I less than 220 microns in length bisetis — 2 antepygidial bristles per side; inner fovea of immovable process exceeds 35 microns; segment V of labial palp exceeds 220 microns in length; crochet spur less than 10 microns in length; finger of clasper exceeds 120 microns in length weaveri 9(7). Ventrolateral lobe of sternum VIII divided into upper convex lobe and lower acuminate lobe; crochet spur less than 10 microns in length 10 — Ventrolateral lobe of sternum VIII not divided; crochet spur exceeds 10 microns in length I ' 10(9). Width of median dorsal lobe of aedeagus exceeds 20 microns; segment V of labial palp exceeds 140 microns in length; depth of sinus in sternum VIII less than 45 microns procus procus — Width of median dorsal lobe of aedeagus less than 20 microns; segment V of labial palp less than 130 microns in length; depth of sinus in sternum VIII exceeds 45 microns procus muiri 11(9). Inner fovea of immovable process less than 20 microns below the dorsal mar- gin; crochet spur less than 20 microns in length; sternum IX less than 180 microns in length cavernicolus 376 Great Basin Naturalist Vol. 39, No. 4 — Inner fovea of immovable process more than 25 microns from dorsal margin; crochet spur exceeds 30 microns in length; sternum IX exceeds 180 microns in length 12 12(11). Inner fovea of immovable process less than 20 microns in length; crochet spur less than 60 microns in length; segment V of labial palp more than 170 microns in length divisus — Inner fovea of immovable process exceeds 40 microns in length; crochet spur exceeds 60 microns in length; segment V of labial palp less than 170 microns in length wilsoni Key to Species of Megaiihroglossiis (Females) 1. Bulga of spermatheca compressed (i.e., appears to be withdrawn into itself (Fig. 86) 5 — Bulga of spermatheca not compressed (Fig. 84) 2 2(1). Posterior margin of sternum VII with sinus 3 — Posterior margin of sternum VII without sinus 4 3(2). With 3 antepygidial bristles per side; segment V of labial palp less than 300 microns in length; hilla of spermatheca less than 40 microns in width (Fig. 94) . spenceri — With 2 antepygidial bristles per side; segment V of labial palp exceeds 300 microns in length; hilla of spermatheca more than 40 microns in width weaveri 4(2). With 3 or 4 antepygidial bristles per side; pronotal comb with 7-8 spines per side; segment V of labial palp less than 230 microns in length; hilla of spermatheca less than 40 microns in width cavernicolus — With 3 antepygidial bristles per side; pronotal comb with 8-9 spines per side; segment V of labial palp more than 270 microns in length; hilla of spermatheca more than 40 microns in width jamesoni 5(1). Posterior margin of sternum VII with sinus, more than 10 microns in depth (Fig. 33) 8 — Posterior margin of sternum VII usually without sinus but if present then less than 10 microns in depth 6 6(5). Distal portion of hilla of spermatheca more than 115 microns, more than 50 microns in width; metatarsal segment I more than 300 microns in length sienae — Distal portion of hilla of spermatheca less than 114 microns, less than 50 microns in width; metatarsal segment I less than 290 microns in length 7 7(6). With 3 antepygidial bristles; pronotal comb with 6 spines per side; sperina- thecal hilla ratio (vertical length: horizontal length) exceeds 1.50; spermathecal bulga ratio (width of bulga at point of greatest width; width of bulga at point of narrowest width) exceeds 1.50. (Fig. 34) procus muiri — With 2-3 antepygidial bristles: pronotal comb with 6-8 spines per side; spermathecal hilla ratio less than 1.20; spermathecal bulga ratio less than 1.40 procus procus 8(5). Hilla of spermatheca more than 40 microns in width; sternum VII sinus depth to width ratio less than 2.00; with 2 antepygidial bristles per side; pronotal comb with 8 spines per side; spermathecal hilla ratio less than 1.10 with bulga ratio exceeds 1.50 bisetis December 1979 Tipton et al.: Anomiopsyllinae 377 — Hilla of spermatheca less than 40 microns in width; sternum VII sinus depth to width ratio exceeds 2.(K); antepygidial bristles variable, 1-4 per side; pronotal comb spines variable, 6-8 per side; spermathecal hilla exceeds 1.10 with bulj^a ratio exceeds 1.70 (if hilla ratio less than 1.10, then bulga ratio less than 1.50) ... ! 9 9(8). Sternum VII sinus depth to width ratio exceeds 2.40; with 2-4 antepyi^idial bristles per side; pronotal comb with 6-8 spines per side; segment V of labial palp exceeds 270 microns in length (average) 10 — Sternum VII sinus depth to width ratio less than 2.40; with 3 antepygidial bristles per side; pronotal comb with 7-8 spines per side; segment V of labial palp less than 270 microns in length (average) 11 10(9). Sternum VII sinus ratio exceeds 4.00; spermathecal hilla ratio exceeds 1.20 and bulga ratio exceeds 1.80; pronotal comb with 8 spines per side siniti — Sternum VII sinus ratio less than 3.00; spermathecal hilla ratio less than 1.20 and bulga ratio less than 1.70; pronotal comb with 6-8 spines per side divisus 11(9). Pronotal comb with 8 spines per side; spermathecal hilla width exceeds 35 mi- crons; segment V of labial palp more than 260 microns in length; spermathecal hilla ratio less than 1.10 and bulga ratio less than 1.50 becki — Pronotal comb with 7 spines per side; spermathecal hilla width less than 35 microns; segment V of labial palp less than 260 microns in length; spermathecal hilla ratio exceeds 1.20 and bulga ratio exceeds 1.80 wilsoni Mendez (1956) pointed out that many of the morphological characters of the female are variable and unreliable; therefore, he did not include a key to the females in his paper. It is important to consider the following when using the above key for females: (1) Definitive determination of the female should be based on presence of the male. (2) The key is useful if only females are collect- ed. (3) The key does not include Megarthro- glossiis sicarnns because a specimen was not available to us; however, it would occur in couplet 4 along with M. cavernicolus and M. iamesoni. Previous authors have used the phrase (when referring to the posterior margin of sternum VII) "sinus lacking," or "sinus pres- ent but shallow." In many instances, the dif- ference between lacking a sinus to having a shallow sinus is only an interpretation of the author. To clarify this point, we have used a -comparison of depth to width. A width to depth ratio that exceeds 5 means that the margin has only a slight concavity and ap- pears to lack a well-defined sinus. Megarthroglossiis becki Tipton & Allred Figs. 35, 37, 40, 42, 68, 82, 10() Megarthwglossus hecki Tipton and Allred 1951:108, 113; Mendez 1956:166; Stark 1959:100; Parker and How- ell 1959:597-604; Hopkins and Rothschild 1962:380; Jel- lison and Glesne 1967:167; Tipton and Saunders 1971:18; Mendez and Haas 1972:285-288; Mendez and Haas 1973:1132; Lewis 1974:15.5. Type host.— Neotoma cinerea acmia Nest. Type locality.— Buckley's Mine, Rock Canyon, Provo, Utah Co., Utah. Type specimens.— United States National Museum, Washington, D.C. Diagnosis.- Male: The ventrolateral lobe of sternum VIII has a long, fingerlike cau- dally curved extension which distinguishes it from all other species (Fig. 40). The inner fovea of the immovable process (Fig. 42) of the clasper is located on the caudal margin and is more than 50 microns below dorsal margin of clasper. The hump on the dorsal margin of the aedeagus varies in height from 4.4 to 11.0 microns (average of 8.0). In most of the specimens measured, the hump was observed as only a slight swelling in the area 378 Great Basin Naturalist Vol. 39, No. 4 Fig. 32. Measurements which show the position of the inner fovea in relation to the dorsal margin of the immovable process of the clasper. A represents the dis- tance of the fovea from the dorsal margin of the clasper. B represents the length of the inner fovea. The inner fovea in this illustration is submarginal. Fig. .33. Measurements of the sinus in the caudal mar gin of the female sternum VII. A represents the depth o the sinus and B represents the width of the sinus. Tht sinus ratio is equal to B divided by A. Fig. 34. Measurements of the spermatheca: A over B = hilla ratio; C over D = bulga ratio; E = width of hiUa. of the dorsal margin between the crescent sclerite and the sclerotized inner tube (S.I.T.). Female: The sinus in the posterior margin of sternimi VII is more than 10 microns in depth or the depth to width ratio is less than 2.40 (see Fig. 2). Segment V of the labial palp is less than 270 microns in length (average). Megarthroglossus becki can be separated from M. wilsoni, its closest relative, in that the hilla is more than 35 microns in width, and it is less than 35 microns in M. wilsoni. Distribution.— Utah: Kane, Piute, Utah, and Wayne counties. Material examined.— Utah: (Kane Co.) 5$, 3$, ex Neotoma albigida, 5-XII-1971, BYU; 1 (5 , ex Neotoma cinerea, 13-X-1971, BYU; 1 (5 , ex Peromijscus crinitiis, 11-XII- 1971, BYU (these three collections were made i .. the NAV-KAI project site); (Piute Co.) 1 $ , ex Thomomijs bottae, Kingston, 26- VI- 1952, Killpack and Beck, BYU; (Utah Co.) 3^,7? paratypes, ex Neotoma cinerea nest, Buckley's Mine, Provo, 24-XI-1949, Allred, BYU; 4 ? , same host, same location 21-X- 1950, Allred, BYU; 4 <5 , 4 ? , same host, Pro- vo, 25-XI-1948, Tipton, BYU; 1 ? , same host, Rock Canyon, Provo, 24-11-1951, Allred and Beck, BYU; 1 ? , same host, same location, 30-III-1951. Allred and Beck, BYU; 2(? , 3 ? , ex Neotoma cinerea. Aspen Grove, American Fork Canyon, American Fork, 13-X-1951, Bamum et al., BYU; 1$ , 19 $ , same host, Thistle, 2-XI-1951, Barnum et al., BYU; 3 ? , same host, Spanish Fork Canvon, Spanish Fork, 2-XI-1951, Barnum et al., BYU. Discussion.— This species is associated with Neotoma cinerea ssp. and more particu- larly with the nests of this host. Neotoma cin- erea occurs in Utah, Idaho, Arizona, Nevada, and California; however, M. becki has been collected only in Utah. It is likely that it has a broader distribution than collection records indicate. Megarthroglossus bisetis Jordan & Rothschild Figs. 45, 57, 72, 85, 101 Mcgartiiroglossits hisctis Jordan and Rothschild 1915:54; Jellison and Good 1942:83; Ewing and Fox 1943:112; Fads and Menzies 1949:33-39; Fads 1950:54; Traub and Hoff 1951:1-23; Williams and Hoff 1951:310-311; Jellison, Locker, and Bacon 1953:104; Morlan 1954:446-448; Morlan 1955:93-125; Mendez 1956:167-168; Hopkins and Rothschild 1962:390; Jellison December 1979 Tipton et al.: Anomiopsyllinae 379 and Glesue 1967:168; Rail et al. 1969:92-94; Forcuni et al. 1969:412; Miller et al. 1970:698, 7(K)-7()1; Clark et al. 1971:1191; Mendez and Haas 1972:285-288; Mendez and Haas 1973:11.32; Lewis 1974:155. \h-g,(irthw^lossus divisus bisetis: Hubbard 1947:302; Jellison and Senger 1976:79. Type host.— Neotoma sp. Type locality.— Beulah, San Miguel Co., New Mexico. Type specimens.— British Museum of Nat- ural History, South Kensington, London, England. Diagnosis.— Male: The posterior margin of sternum VIII is evenly convex and there are 2 or 3 (usually 2) antepygidial bristles as in M. weaveri. Megarthroglossus bisetis may be separated from M. weaveri on the basis of the following characters: the inner fovea of the immovable process is less than 35 mi- crons from the dorsal margin; segment V of labial palp is less than 220 microns in length; the spur of the crochet is more than 10 mi- crons in length; and the finger of the clasper is less than 120 microns in length. Female: Megarthroglossus bisetis, M. weaveri, and some specimens of M. siniti and M. divisus possess two antepygidial bristles. Megarthroglossus bisetis may be separated from M. smiti and M. divisus in that the hilla of the spermatheca exceeds 30 microns in width and sternum VII has a sinus depth-to- width ratio of less than 2.00, whereas this ra- tio is greater than 2.00 in M. smiti and M. di- visus. Megarthroglossus bisetis can be sepa- rated from M. weaveri in that the former has a bulga which is compressed (the portion of the bulga to which the spermathecal duct at- taches appears to be withdrawn or pu.shed into the other portion of the bulg-a). Fig. .35. Megarthroglossus hecki: Male, head and thorax including coxae. 380 Great Basin Naturalist Vol. 39, No. 4 Figs. 36-38. Males, metathoracic legs; 36, Cdllistopsiilhs tcrinus dcutcnis- 37. Mcalifornia, l.os Angeles, California. Diagnosis.— Male: The two subspecies of MegarthroglosHUfi proem may be separated from all other species of Megarthroiijossiis by the following characters: The ventrc^lateral F.gs. 43-46. Males, claspers; 43, Megarthroglossus .ilsonU 44, M. .icornus. 45, M. hiseti. 46, M. .eaveri. 386 Great Basin Naturalist Vol. 39, No. 4 lobe of sternum VIII is divided into an upper evenly convex lobe and a lower acuminate lobe separated by a sharp sinus; the crochet does not possess the well-defined spur ob- served in other species such as M. divisiis; however, a short lobe less than 10 microns may be present in some. Megarthroglossus p. miiiri can be distinguished from M. p. procus in that the latter has a median dorsal lobe which is less than 20 microns in width; seg- ment V of the labial palp is 130 microns in length and the sinus in sternum VIII exceeds 45 microns in depth. Female: Usually there is no sinus in the Figs. 47-50. Males, claspers; 47. Me^orthroolossus divisus (Wayne Co.. Utali); 48. M. divisus (Ravall. Co., Mon- tana); 49, M. procus muiii; 50, M. p. procus. December 1979 Tipton et al.: Anomiopsyllinae 387 posterior margin of sternum VII, but, if pres- ent, it is less than 10 microns in depth; there are 3 antepygidial bristles per side; the pro- notal comb has 6 spines per side; the hilla ra- tio exceeds 1.50, and the bulga ratio exceeds 1.50 (Fig. 34). These characters are adequate for separation of M. p. procus and M. p. miiiri, but the presence of males in a collec- tion aids greatly in subspecific discrimina- tion. Distribution.— California: Fresno, Mono, and Plumas counties. Material examined.— California: (Fresno Co.) 1 S , holotype, ex Tamiasciunis douglasi olbolimbatus. Tally's Hole, 25-VIII- 1941, Auguston; 1 $ , allotype, ex Sorex sp. navigator, Tully's Hole, 24-VIII-1941, Au- gustson; (Plumas Co.) 1 $ , ex Peromyscus maniculatus 6.4 km E of Quincy, 1520 m, 24- X-1949, Jameson. Host synonymy.— Tamiasciunis douglasi albolimbatus = Tamiasciurus hudsonicus al- bolimbatus. Discussion.— Megarthroglossus procus muiri has been collected from species of Ta- miasciurus, Ochotona, and Sorex. Additional specimens are necessary to determine host as- sociations of this flea. It has been collected only in California; however, it is likely that it occurs in Nevada, Oregon, and Washington as well. Megarthroglossus procus procus Jordan & Rothschild Figs. 50, 62, 76, 77, 88, 89, 101 Megarthroglossus procus Jordan and Rothschild 1915:47-50; Wagner 1930:130; Spencer 1936:14; Mail and Holland 1940:126; Hubbard 1940:37(4); Augustson 1941:151; Ewingand Fox 1943:113; Hubbard 1943:1-12; Hatt 1943:311-345; Hubbard 1947:297; Hubbard 1949: 126; Holland 1949a: 10; Holland 1949b; 100; Tipton 1950:63; Hopkins 1952:363-365; Mendez 1956:164-166; Jameson and Brennan 1957:45-54; Stark 1959:100; Beck 1966:76; Beck and Ailred 1966:13; Jellison and Glesne 1967:172; Tipton and Saunders 1971:18; Mendez and Haas 1972:285-288; Mendez and Haas 1973:1132; Lewis 1974:155; Egoscue 1976:478; Jellison and Sanger 1976:80. Megarthroglossus similis Wagner 1936:196; Mail and Holland 1939:126; Jellison and Good 1942:84; Ewing and Fox 1943:113; Hubbard 1943:1-12; Hubbard 1947: 302; Holland 1949a: 10; Holland 1949b:99; Jellison, Lock- er, and Bacon 1953;107; Mendez 19.56:175. MegartJirogJossus senislcs (sic) Spencer 1936:14. Megarthroglossus procus oregonesis Hubbard 1947: 299-300; Jellison, Locker, and Bacon 1953:105. Megarthroglossus procus procus: Hubbard 1947:297; Jellison. Locker, and Bacon 1953:105; Hopkins and Hothschild 1962:.37 1-372. Type host.— Spilogale sp. Type locality.— Chilliwack, British Co- lumbia, Canada. Type specimens.- British Must-urn (Natu- ral History), London. Diagnosis.— Male: The two subspecies of M. procus have a divided ventrolateral lobe of sternum VIII which is not present in other species of the genus Megarthroglossus. M. p. procus may be separated from M. p. muiri in that in the former the median dorsal lobe of the aedeagus is more than 20 microns in width; segment V of the labial palp is more than 140 microns in length; and the sinus in sternum VIII is less than 45 microns in depth. Female: The pronotal comb has 6-8 spines per side; there are 2 or 3 antepvgidial bristles; there is no sinus in the caudal margin of sternum VII or, if it is present, it is less than 10 microns in depth; the hilla ratio is less than 1.20 and the bulga ratio is less than 1.40. However, a definite identification is possible only if male specimens are available. Distribution.— California: Eldorado, Plumas, and San Bernardino counties. Colorado: Montezuma County. Nebraska: Sioux County. Nevada: Douglas and Washoe counties. Oregon: Hood River, Linn, and Washington counties. Washington: Skagit, Whatcom, and Yakima counties. Wyoming: Laramie and Weston counties. Utah: Utah County. Canada: Briti.sh Columbia. Material examined.— California: (El Dorado Co.) 2 males, 1 female, ex chipmunk, no specific location, X-1936, Eskey (Jellison); (San Bernardino Co.) 1 male, ex Citellus lat- eralis. Big Bear Lake, 13-V-1955, Martin. Nevada: (Washoe Co.) 1 S , I $ , ex chip- munk, no specific location, X-1936, Eskey (Jellison). Oregon: (Linn Co.) 1 ? , ex Ta- miasciurus douglasi, 2.4 km N of Big Lake, 1- XI-1969, Maser; 1 $ , ex Eutamias towm- endi, T14S, R6E, NE V4, Sec. 28, 1400 m, 12- IX-1972, Maser; 1 ? , ex Peromyscus manicu- latus, same location, same elevation, same date, Maser; 1 ? , ex Glaucomys sabrinus, T15S R15E, SEV4, Sec. 11, 26-IX-1972, Ma- ser; 1 $ , same host, T25S, R6E, SE'/4, Sec. 11, 1040 m, same date, Maser; 1 $ , same host, T15S, R5E, SO 1/4, Sec. 11, same eleva- 388 Great Basin Naturalist Vol. 39, No. 4 tion, 28-IX-1972, Maser; (Washington Co.) 1 (5,1$, paratypes, ex Tamiasciurus d. dotig- lasi, Caston, 7-III-1932, Hubbard (BYU); 1 5,1$, paratypes same host, same location, 8-III-1932, Hubbard (BYU). Washington: (Skagit Co.) 2 $ , 7 ? , ex Neotoma nest, Blanchard Caves, 19-III-1966, Senger; 3 ? , same host, same location, XII- 1965, Senger; 10 (5 , 12 $ , same host, same location, 20-XI- 1966, Senger; 15 (5 , 20 $ , same host, same location, 5-XI-1967, Senger; 1 ? , same host, same location, 18-IX-1967, Senger; 3 5,5 ? , same host, same location, 17-XI-1968, Senger; 3 $ , 2 ? , same host, same location, 9-III-1968, Senger; 6 <5 , 17 ? , same host, same location, XI- 1965, Senger; 15 S , 5 ? , Figs. 51-54. Males, claspers; 51, Megarthmghssm smith 52, M. spencerh 53, M. siemie: .54, .\/. jamcsoni. December 1979 Tipton et al.: Anomiopsyllinae 389 55 Figs. 55-58. Males, sterna VIII and IX; 55, Me,ar,k,o,lossus .UsonU 56, M. sica.u,s. 57. A/, ^.is.is, 58, M. weaveri. 390 Great Basin Naturalist Vol. 39, No. 4 same host, same location, 20-XI-1965, Seng- er; 1 ? , same host, same location, III- 1965, Senger; 24 S , 21 ? , same host, Lizzard Caves, 19-11-1966, Senger; 2 $ , 6 ? , same host, same location, 15-1-1966, Senger; 2 $ , 1 ? , same host, same location, XI- 1965, Senger; (Wheaton Co.) 5 $ ,5 ? , same host. Glacier, X-1963, Senger; 1 ? , ex nest in old fin snag, Chuckanut Mt., 14-11-1965, Senger; 1 ? , ex nest in fallen snag, Silver-Fir Camp- ground, Nooksack River, 3-IX-1966, Senger; (Yakima Co.) 2 ? , ex Ochotona princeps, Chinook Pass, 16-IX-1966, Senger; Canada: (British Columbia) 1 S , I ? , ex Tamias- ciimis douglasi, 11.2 km S of Boston Bar, 14- III-1948, Holland; I S , ex Glaucomys sp.. Grouse Mt., Vancouver, 21-1-1947, Dowding (Holland); 1 (5 , ex Sciunis douglasi, Gambien Island, 21-11-1943, Holland; 1 ? , ex squirrel, Cultus Lake, 31-X-1947, Leavens (Holland); 1 ? , ex Tamiasciurus sp., Huntingdon, 16- XI-1946, Racey (Holland); 1 ? , ex Rattus norvegicus, Vancouver, 7-1-1945, Holland. Host synonymy.— Tamiasckirus d. doug- lasi = Tamiasciurus hudsoniciis douglasi; Sciurus douglasi = Tamiasciurus hudsonicus douglasi; Citellus lateralis = Spermophilus lateralis. Discussion.— Most of the specimens in Dr. C. M. Senger's collection were taken from species of Neotoma in Washington. However, it has been collected from several other hosts, principally, Tamiasciurus hudso- nicus douglasi. Diagnosis.- Male: The caudal margin of sternum VIII is evenly convex in M. sicamus, M. bisetis and M. iveaveri. However, M. si- camus has a hump on the dorsal margin of the aedeagus that is more than 30 microns while it is less than 10 microns in the other two species. Other species which have a hump on the dorsal margin of the aedeagus include M. spenceri, M. smiti, M. sierrae, M. iamesoni, and M. hecki. The inner fovea of the immovable process of the clasper is more than 60 microns below the dorsal margin in M. sicamus but not in the other five species mentioned. Female: No specimens were available for study. On the basis of the literature, it ap- pears that M. sicamus most closely resembles M. cavernicolus and M. jarnesoni. The hilla appears to be less than 40 microns in width but more than 40 microns in M. jamesoni. In M. sicamus the hilla is bent toward the bulga but in M. cavernicolus it is not. Distribution.— Canada: British Colum- bia. Material examined.— Canada: (British Columbia) 1 ? , ex Lynx sp., Kamloops, 20- V-1946, Carter (Holland); 1 ? , ex Neotoma, Pavilion Lake, Pavilion, 5-VII-1950, Holland; \ S ,ex Canis latrans. Eagle River, Sicamous, 6-IX-1903, Dippie (Smit). Discussion.— Megatihroglossus sicamus has been collected only from British Colum- bia. Because of the paucity of specimens, re- liable host association data are lacking. Megarthroglossus sicamus Jordan & Rothschild Figs. 44, 56, 81,91, 101 Megarthroglossus sicamus Jordan and Rothschild 1915:50-52; Dalla Torre 1924;17; Wagner 1936;196; Mail and Holland 1939:126; Jellison and Good 1942:84; Ewing and Fox 1943:113; Hubbard 1943:1-12; Costa Lima and Hathaway 1946:125; Hubbard 1947:302; Hol- land 1949a: 10; Holland 1949b; 101; Jellison, Locker and Bacon 1953:104; Mendez 1956:173-174; Hopkins and Rothschild 1962:382; Lewis 1974:155. Type host.— Canis latrans. Type locality.— Eagle River, Sicamous, British Columbia, Canada. Type specimens.— British Museum (Natu- ral History), South Kensington, London, Eng- land. Megarthroglossus sierrae Auguston Figs. 53, 65. 69, 92, KH) ' Megarthroglossus divisus sierrae Augustson 1953: 125-126; Jellison and Glesne 1967:170; Jellison and Senger 1976:79. Megarthroglossus sierrae Mendez 1956:174; Hopkins and Rothschild 1962:378; Lewis 1974:155; Jellison and Senger 1976:80. Type host.— Ochotona schisticeps muiri and Tamiasciurus douglasi albolimbatus. Type locality.- Cascade Valley, Fresno Co., California. Type specimens.— Allan Hancock Founda- tion, University of Southern California, Los Angeles, California. Diagnosis.— Male: Af. sierrae and M. jamesoni resemble each other in that the me- December 1979 Tipton et al.: Anomiopsyi.mnak 391 Figs. 59-62. Males, sterna VIII and IX; 59 Megarihroghssus divisus (Wayne Cc Co., Montana); 61, M. proctis mniri; 62, M. p. proctis. Utah); 60. A/, dhisus (Ravalli 392 Great Basin Naturalist Vol. 39, No. 4 Figs. 63-66. Males, sterna VIII and IX; 6.3, Megarthroglossus spenceri; 64, Xf. smiti; 6,5. ,\/. .siernie; 66, M. jamesoni. December 1979 Tipton et al.: Anomiopsyllinae 393 Figs. 67-69. Males, aedeagus; 67, Megarthroglossus jamesoni; 68, M. becki; 69, M. sierrae. 394 Great Basin Naturalist Vol. 39, No. 4 dian dorsal lobe of the aedeagus is reduced to a short blunt lobe in both species. M. sierme may be distinguished from M. jaiyiesoni on the basis of the following characters: The hump on the dorsal margin of the aedeagus is less than 20 microns; segment V of the labial palp is less than 220 microns in length; the inner fovea of the immovable process is mar- ginal, and the ventrolateral lobe of sternum VIII is subtnincate. Female: M. sierrae and M. procus ssp. each lack a sinus in the caudal margin of sternum VII, but, if a sinus is present, it is less than 10 microns in depth. In M. sierrae the vertical portion of the hilla is more than 115 microns and the hilla is more than 50 microns in width; metatarsal segment I is more than 300 microns in length. Distribution.— California: Fresno and Mono counties. Material examined.— California: (Fresno Co.) 1 S , holotype, ex Ochotona schisticeps muiri. Cascade Valley, 27-VIII- 1941, Augustson; (Mono Co.) 1 ? , allotype, ex Tamiasciuriis douglasi albolimbatits. Mammoth Lakes, 31 -VII- 1939, Augustson. Host synonymy.— Tamiasciiirus douglasi alholimhatus = Tamiasciuriis hudsonicus al- bolirnbatus. Discussion.— Megarthroglossus sierrae has been reported only from the type locality in California on Tamiasciurus and Ochotona. Megarthroglossus sierrae, M. divisus, and M. procus overlap in California in both ecologi- cal and geographical distribution. Additional data is needed to clarify the relationships of these species. Megarthroglossus smiti Mendez Figs. 51, 64, 79, 93, 101 Megarthroglossus smiti Mendez 1956:175; Howell 1955:35-48; Howell 1957:566-573; Parker and Howell 1959:597-604; Stark 1959:99; Hopkins and Rothschild 1962:385; Jellison and Glesne 1967:173; Tipton and Saunders 1971:18; Mendez and Haas 1973:1132; Jenkins and Grundmann 1973:81; Lewis 1974:155; Egoscue 1976:479; Jellison and Senger 1976:80. Type host.— Neotoma lepida nest. Type locality.— Lynndyl, Millard Co., Utah. Type specimens.— United States National Museum, Washington, D.C. Diagnosis.— Male: M. smiti, like M. sierrae and M. jajiiesoni, has a hump on the dorsal margin of the aedeagus; however, it is more pronounced in M. smiti than in M. sierrae and less so than in M. jamesoni. There is a considerable amount of variability in this character and so its taxonomic value is ques- tionable. In M. smiti the median dorsal lobe is produced into a long process which curves caudally at the apex, and sternum VIII has an undulating posterior margin with a short, subacuminate lobe. Female: M. smiti, M. divisus, M. becki, and M. wilsoni are similar in many respects. M. smiti may be distinguished from the latter two species in that the depth to width ratio of the sinus in sternum VII is more than 3.0; segment V of the labial palp is more than 270 microns in length (average). Megarthroglossus smiti may be separated from M. divisus in that the sternum VII sinus ratio is more than 4.0; the hilla ratio exceeds 1.20 and the bulga ratio exceeds 1.80. Distribution.— Nevada: Nye County. Utah: Beaver, Emery, Juab, Millard, Tooele, and Utah counties. Material examined.— Nevada: (Nye Co.) 1 (5 , ex Neotoma lepida. Mercury, 6-XI-1961, AEC-NRTS (BYU). Utah: (Juab Co.) 5 $ ,2 ? , ex Neotoma lepida nest, Jericho, 29-IV- 1954, J. F. Howell; 1 S , I ? , ex A^. lepida 4.5 km N Jericho, 8-V-1954, J. F. Howell; 9 (5 , 16 ? , ex N. lepidci nest, Jericho, 8-X- 1954, J. F. Howell, 5 yllus. How- ever, the latter genus does not have a pro- notal comb. The mesocoxa has an incomplete longitudinal break that extends for less than one-half the diagonal distance. Stenistomera has a comb of dorsolateral bristles on the mesotibia and Callistopsyllus does not. In ad- dition, the bristles of the head are more nu- merous and prominent in Stenistomera than in other genera of the subfamily. Description.— Head: .\nterior margin convex, angulate or helmet shaped; numerous stout (spiniform in alpina) marginal and sub- marginal bristles; pores and placoids scat- tered over frons and occiput, but mostly sub- marginal. Eye absent. Antenna extremely short. Maxillary lobe reduced, angulate. Max- illary palp 4-segmented, short, extends to midpoint on forecoxa. Labial palp well de- veloped, 4-segmented, extends to apex of forecoxa, apex more or less asymmetrical. Thorax: Pronotal comb of 16-18 teeth. Lateral metanotal area absent. Pleural arch absent. Metepisternum fused with metaste- n.im. Hind coxa narrow, false combs on meso- tibiae, four pairs lateral bristles on fifth tarsal segment. Abdomen: One row of bristles on each ab- dominal tergum. Three antepygidial bristles in both sexes. Modified abdominal segments— Male: Fixed process of clasper not well developed; movable process long, narrow, with sides sub- 404 Great Basin Naturalist Vol. 39, No. 4 DISTRIBUTION OF THE GENUS MEGARTHROGLOSSUS ° M. becki * M. cavernicolus • M. a. divisus " M. lamesoni o M. sierrae ■ M. wilsoni Fig. 10(). Distribution of Megarthroglossus speci and M. wilsoni. M. becki, M. cavernicolus, M. (Uiisiis, M. jamesoni, M. sierrae. December 1979 Tipton et al.: Anomiopsyllinae ■\m DISTRIBUTION OF THE GENUS MEGARTHROGLOSSUS • M.bicetis ° M. p. muiri * M. p. procus ♦ M. Sicamus ° M. sm ■ M. spencer ^ M. weaver I spenceri, and M. weaveri. 406 Great Basin Naturalist Vol. 39, No. 4 parallel, apex truncate; with two well-pig- mented, spiniform, apical or subapical setae. Sternum IX with narrow proximal arm but with subapical portion swollen; distal arm ex- panded, bearing subapical spiniform(s) plus several small marginal and submarginal setae. Aedeagus with narrow, saberlike apodeme, with long-coiled appendage; apodemal rods long, coiled. Crescent sclerite and capsule prominent. Median dorsal lobe well sclero- tized, apex somewhat hooklike. Modified abdominal segments— Female: Posterior margin of sternum VII convex dor- sally, with ventral sinus. Bulga of sperma- theca barrel shaped; hilla bent at right angles to bulga. 2(1). Key to the Species of Stenistomera (Male) Process of clasper broader than distal arm of sternum IX at its widest point (Fig. 107); two spiniform bristles on caudal margin separated by more than 3 times length of one spiniform; distal arm of sternum IX with apex subacuminate (Fig. 104) macwdactijla Process of clasper narrower than distal arm of sternum IX at its widest point; two spiniform bristles on caudal margin separated by less than 3 times length of one spiniform, distal arm of sternum IX with apex subacuminate to subtruncate 2 Distal arm of sternum IX with two prominent subapical bristles; apex subacuminate (Fig. 105) hubhardi Distal arm of sternum IX with one prominent subapical bristle; apex subtruncate (Fig. 103) alpina Stenistomera hubhardi Egoscue Figs. 28, 105, 108, 109, 114, 115 Stenistomera hubhardi Egoscue 1968:138-142; Lewis 1974:155. Type host.— Peromijsciis maniculatus ssp. Type locality.— Crane, Hamey Co., Ore- gon. Type specimens.— United States National Museum, Washington, D.C. Diagnosis.— The labial palp extends for three-fourths the length of the procoxa and the maxillary lobe is subacuminate. The pre- antennal bristles are larger and heavier than in S. macrodactijla, but less so than in S. al- pina. Description.— Male: Movable process of clasper elongate, rectangular, curved slightly caudad, distal one-third with two spiniforms on caudal margin. Base of acetabular bristle midway between sensilial plate and articu- lation of movable process of clasper. Distal portion of sternum IX blade shaped, with two thick, blunt, subapical bristles. Female: Three antepygidial bristles ap- proximately equal in length; anal stylet long, about four times longer than wide, with long subapical bristle plus shorter apical bristle. Bulga of spermatheca not markedly dissimilar in shape from that of other members in genus or in CaUistopsijUus terinus; hilla sharply bent (approximately 90 degrees) shortly after exiting from bulga, apex with prominent sclerotized papilla. Distribution.— Oregon: Harney County. Material examined.— Oregon: (Harney Co.) 1 (5 , 1 ? and 1 broken specimen, ex Per- omijscus manicidatus, 8.0 km south of Crane, 23-'X-1966, H. J. Egoscue. Stenistomera macrodoctyla Good Figs. 29, 104, 107, 110, 113, 115 Stenistomera macrodoctyla Good 1943:135; Hubbard 1947:306; Jellison, Locker, and Bacon 1953:186; Hold- enried and Morlan 1955:133-137; Morlan 1955:93-125 Wiseman 1955:1-18; Parker and Howell 1959:597-604 Hopkins and Rothschild 1962:353-354; Beck 1966:77 Jellison and Glesne 1969:302; Allred 1968:78; Egoscue 1968:140; Tipton and Saunders 1971:18; Haas et al. 1973:282; Pratt and Stark 1973:11; Lewis 1974:155; Ego- scue 1976:476. Miochaeta macrodactijla Stark 1959:104. Type host.— Peromijscus eremicus. Type locality.— Mojave Co., Arizona. December 1979 Tipton et al.: Anomiopsyi.linae 407 Type specimens — United States Public Health Service Plague Laboratory, San Fran- cisco, California. Diagnosis.— Labial palp extends to apex of the procoxa. Maxillary lobe is sub- acuminate, but broader at the apex than in S. luihhardi. The preantennal bristles are not enlarged or heavily pigmented as in S. hub- bardi or S. (ilpina. Table 7. Distribution of species of Stenistomeni. Are; Arizona Colorado Idaho Montana Nevada New Mexico Oregon Utah Wyoming British CoKunbia X X X X X X X X X X X X X X X X X Table 8. Host associations of the genus Steimtomera. Host species AmmospennopJiihis h'ticunis Lynx rufiis paUcsccns \eotoiiui alhigula S'eotoinci albigula alhiguki Xeotoma cinerea Neotomci cinerea cieraia Neotoiud einerea dllicolii Neotoma lepida Neotoma lepida lepida Neotoma lepida nevadensi.s Xeotoma niexicana Peromysciis criyiittis Peromyscus crinitus pergracilis Peromysciis tnaiiicidatits Peromyscus manicuhitus sonoriensis Reithrodontomys megalotis megalotis Urocyon cinereoargenteus Spilogale putorius sayatilis Desc:ription.— Male: Movable process of clasper elongate, urn shaped, not curved cau- dad as in S. huhbmdi or S. alpina, with two widely separated spiniforms on caudoniesal surface. Base of acetabvilar bristle near artic- ulation of movable process of clasper. Distal arm of sternum IX narrow at apex, with single thick, blimt subapical bristle. Female: Three antepygidial bristles of about equal length. Anal stylet long, (about three to four times as long as wide), with row of four ventral marginal bristles of about equal length (variable, usually two to five), one long subapical bristle, plus one short apical bristle. Caudal margin of sternum VII with dorsal portion evenly convex, short sub- acuminate to triangular ventral lobe ventral to broad, shallow sinus. Bulga tvpically shaped; hilla evenly curved dorsad, lacks sclerotized papilla as in S. hubhardi. Distribution.— Colorado: Montezuma County. Idaho: Bingham and Butte counties. Nev.\da: Washoe County. New Mexico: Sandoval County. Utah: Daggett, Iron, San Juan, Tooele, and Uintah counties. Wyoming: Sweetwater County. Material examined.— Colorado: (Mon- tezuma Co.) 1 5,3 ? , ex Peromyscus ma- niculatus, Mesa Verde National Park, 26-V- 1962, C. Douglas (BYU). Idaho: (Bingham Co.) 1 ? , ex Peromi/scus manicxdatus, AEC- NRTS (19Y), 22-IX-1966 (BYU); 3 ? , .same host, AEC-NRTS (21-S), 18-IX-1966 (BYU); 1 ? , same host AEC-NRTS (2 IS), 2()-XI-1966 (BYU); 1 5,2 ? , same host, AEC-NRTS (33Y), 17-XI-1967 (BYU); 1 $,\ $, ex ro- dent nest, same location, same date (B\TJ); 14 5,5 ? , ex Pewmi/scus tnaniculotus, AEC- NRTS (33Y), 19X1-1967 (BYU); 1 5 , ex Xeo- toma cinerea, AEC-NRTS (33Y), 22-VIII- 1967 (BYU); (Butte Co.) 3 $ , ex Peromyscus maniculatus, AEC-NRTS (31Y), 16-1-1967 (BYU); 1 S , same host, AEC-NRTS (32Y) 17- XH967 (BYU). Utah: (Daggett Co.) 15,1 2 , ex Peromyscus maniculatus, Linwood, 14- VII- 1954, C. L. Hayward (BYU); (Iron Co.) 1 5 , same host, Parowan, 4-IX-1951, Beck and Allred (BYU); (San Juan Co.) 2 $ , .same host, Bluff, 5-V-1951, D. E. Beck (BYU); 1 $ , same host, Montezuma Creek, 7-VI-1955, Beck et al. (BYU); (Uintah Co.) 1 5,1 ? , ex Peromyscus crinitus, Jensen, 8-XI-1952, Beck and Beck (BYU). 408 Great Basin Naturalist Vol. 39, No. 4 Stenistomera alpina (Baker) Figs. .30, 38, 102. 10.3, 106, HI, 112, 115 TijphlopsijUa alpina Baker 1895:189, 191. Ctenopsylla alpina Wagner 1898:577-578. CtenopsyUus alpinns Baker 1904:427, 452; Triaboschi 1904:285. Stenistomera alpina Rothschild 1915:307; C. Fox 1925:127; Wagner 1930:144; Stanford 1931:153; Wagner 19.39:.32; Eskev and Haas 1940:29, 74; Good 1942:133 JelHson and Good 1942:132; Ewing and Fox 1943:73 Hubbard 1943:1-12; JeUison. Khols. and Mills 1943 1-22; Stanford 1944:175; Hubbard 1947:305; Hubbard 1949:121; Tipton 1950:65; Traub and Hoff 1951:23 pp.; Holland 1952:65-73; Wehrle 1953:37-41; JeUison. Lock- er, and Bacon 1953:186-187; Augustson 1955:36-39; Morlan 1955:93-125; Wiseman 1955:1-23; Holdenried and Morlan 19.56:369-381; Finley 1958:21.3-552; Parker and Howell 1959:597-604; Stark' 1959: 103; Hopkins and Rothschild 1962:350-353; Beck and Allred 1966:13; JeUi- son and Glesne 1967:300-.301; Allred 1968:78; Egoscue 1968:140; Tipton and Saunders 1971:18; Pratt and Stark 1973:11: Haas et al. 1973:282; JeUison and Senger 1973:71; Lewis 1974:155; Egoscue 1976:47.5-488. Delotelis moliatensis Augustson 1942:138; .\uguston 1943:86. Type host.— Moimtain rat (probably Seo- toina cinerea). Type loc.\lity.— Georgetown, Clear Creek Co., Colorado. Type specimens.— United States National Museum, Washington, D.C. Di.\GNOsis.— Labial palp extends bevond the apex of the procoxa and usually bevond the trochanter; the niaxillary lobe is acumi- nate and the preantennal bristles are thick and heavily pigmented; some are spin- iformlike. Description.— Male: Movable process of clasper elongate, curved slightly caudad, with two closely spaced spiniforms on meso- caudal margin, similar to that of S. hubbardi. Base of acetabular bristle displaced toward sensilial plate as in S. Juibbardi, but less so than in S. macrodactyla. Distal arm of ster- num IX somewhat similar to S. hubbardi, but with single, thick, blunt bristle to caudovent- ral angle, apex subtruncate. Female: Three antepygidial bristles, middle bristle longest. Anal stylet long, three to four times as long as wide, usually with two ventral bristles, one long subapical bristle plus short apical bristle. Bulga typical, but with a slight narrowing at end near hilla; no sclerotized papilla on hilla. Distribution.— Arizona: Apache, Coch- ise, and Coconino coimties. Colorado: Clear Creek and El Paso comities. Id.\ho: Butte Countv. Mont.4Na: Custer, Madison, and Ra- Fig. 102. Stenistoviera alpina (Baker): Male, head and thora.x including coxae. December 1979 Tipton et al.: Anomiopsyllinae 409 valli counties. Nevada: Nye and Washoe counties. New Mexico: Chaves, BernaHllo, and Sandoval counties. Oregon: Harnev County. Utah: Cache, Kane, Juab, Salt Lake, San Juan, Sanpete, Sevier, Tooele, and Utah counties. Wyoming: Albany and Sweetwater counties. Canada: Alberta. Material examined.— Arizona: (Cochise Co.) 1 ? , ex Neotoma mexicana, 0.8 km E Buena Vista Park., Chiricahua Mts., 23-XI- 1957, C. E. Ordway (BYU). Idaho: (Butte Co.) 7 (5,5 ? , ex Neotoma cinerea, AEC- NRTS (29), 15-XII-1966 (BYU). Nevada: (Nye Co.) 1 $ , 3 ? , ex Peromysciis crinittis. Mercury, AEC-NRTS, 6-XI1-1961 (BYU); 1 (5,5 $ , ex Neotoma lepida, same location, ll-XII-1960 (BYU); 2 S, n ?, same host, same location, 9-X11-1961 (BYU); 6 (5 , 19 ? , same host, same location, ll-Xll-1961 (BYU); 1 (5,3 ? , same host, same location, 8-XI- 1961 (BYU); 2 (5,2 ? , ex Neotoma albigida, same location, 28-X-1961 (BYU); 13 <5 , 35 ? , ex Neotoma lepida, same location, 6-XII- 1961 (BYU); 1 (5 , ex Ammospermophiliis leucurus, same location, 7-XII-1961; 1 ? , ex Neotoma lepida, same location, 16-XI-1961 (BYU); 1 S , 2 ? , same host, same location, ll-XI-1961 (BYU); 22 $,31 ? , same host, same location, 17-XII-1961 (BYU); 1 ,5,2 ? , same host, same location, 5-1-1962 (BYU); 4 (5,11 ? , same host, same location, 20-Xll- 1961 (BYU); 1 S , same host, same location, 7-1-1962 (BYU); 1 S , same host, same loca- tion, 2-1-1962 (BYU); 1 $ , same host, same location, 2-1-1962 (BYU); 7 5 , 13 ? , same host, same location, 17-XII-1961 (BYU); 1 ,5 , 1 ? , ex Neotoma lepida lepida, Beatty, 27- XII- 1947, C. A. Hubbard (BYU); 27-XII-1947, C. A. Hubbard (BYU). New Mexico: (Ber- nalillo Co.) 1 (5,1 ? , ex Neotoma a. albi- gida, 28.8 km E Albuquerque, 6-III-1949, H. H. Lewis (BYU). Utah: (Kane Co.) 1 $ ,2 ? , ex Urocyon cinereoargenteus, NAV-KAI, 14- XI- 1971 (BYU); 1 ? , ex Peromysciis crinitus, same location, ll-XII-1971 (BYU); (Juab Co.) 8 $ ,2A ? , ex Neotoma lepida lepida. Topaz Mt., 19-1-1964 (Egoscue); 5 $ , \2 ? , same host, same location, 20-1-1964 (Egoscue); 4 5 , 13 ? , same host. Fish Springs, l-XII-1964 (Egoscue); 1 5 , ex Reithrodontomys mega- lotis megalotis, same location, l-XII-1964 (Egoscue); 1 ? , ex Peromysciis maniculatus sonoriensis, Callao, 26-1-1965 (Egoscue); (San Juan Co.) 2 ,5 , ex Neotoma cinerea, S Moab, 8-V-1951, Allred and Myklebust (BYU); (Tooele Co.) 2 ^ , 1 $ , ex Lynx rufus palles- cens, 9.6 km E Granite Mt., 1.300 m. 5-III- 1965, H. J. Egoscue (Egoscue); 1 <5 , ex Neo- toma lepida lepida, granite outcrops, E Gran- ite Mt., 1380 m 19-XI-1965, H. J. Egoscue (Egoscue); 5 $ , same host, same location, same elevation, 13-1-1966, H. J. Egoscue (Egoscue); 2 (5,5 ? , same host, same loca- tion, same elevation, 18-1-1966, H. J. Egoscue (Egoscue); 5 $ , 8 ? , same host, same loca- tion, same elevation, 24-III-1966, H. J. Ego- scue (Egoscue); 6 $ , same host, same loca- tion, same elevation, 9-III-1966, H. J. Egoscue (Egoscue); 3 $ , same host, same lo- cation, same elevation, 15-III-1966; H. J. Egoscue (Egoscue); 4 ? , 2 ,5 , same host, same location, same elevation, 25-11-1966, H. J. Egoscue (Egoscue); 2 $ , same host, same location, same elevation, 2-III-1966; H. J. Egoscue (Egoscue); 1 $ , same host, same lo- cation, same elevation, 19-XI-1965, H. J. Egoscue (Egoscue); 1 $ , same host, same lo- cation, same elevation, 19-11-1966, H. J. Ego- scue (Egoscue); 1 $ , 3 $ , same host, same location, same elevation, 17-11-1966, H. J. Egoscue (Egoscue); 1 $ , I ? , same host, same location, same elevation, 17-XI-1965, H. J. Egoscue (Egoscue); 1 $ , 3 $ , same host, same location, same elevation, 13-1- 1966, H. J. Egoscue (Egoscue); 2 $, 1 $ , ex Peromysciis crinitus pergracilis, same loca- tion, same elevation, 12-1-1966, H. J. Egoscue (Egoscue); 1 S , same host, same location, same elevation, 16-III-1966, H. J. Egoscue (Egoscue); 1 $ , same host, same location, same elevation, 18-1-1966, H. J. Egoscue (Egoscue); 1 ? , same host, same location, same elevation, 11-1-1966, H. J. Egoscue (Egoscue); 1 (5,2 ? , ex Spilogale ptitorius sayatilis, same location, same elevation, 19-1- 1966, H. J. Egoscue (Egoscue); 1 (5 , 6 $ . ex Peromysciis crinitus pergracilis. Little Granite Mt., 1440 m, 6-XII-1961, H. J. Egoscue (Ego- scue); 1 (5,2 ? , ex Neotoma lepida lepida, same location, same elevation, 6-XII-1966, H. J. Egoscue (Egoscue); 1 S , 2 $ , same host, same location, same elevation, 8-XII-1966, H. J. Egoscue (Egoscue); 1 S , I $ , same host, same location, same elevation, 14-XII-1966, H. J. Egoscue (Egoscue); 8 $ , 7 $ , same host, same location, same elevation, 15-XII- 410 Great Basin Naturalist Vol. 39, No. 4 1966, H. J. Egoscue (Egoscue); 1 $ , 2 ? , same host, same location, same elevation, 20- XII- 1966, H. J. Egoscue (Egoscue); 4^,4?, same host, same location, same elevation, 21- XII- 1966, H. J. Egoscue (Egoscue); 4 <5 , 10 ? , same host, same location, same elevation, 6-1-1967, H. J. Egoscue (Egoscue); 5 S ,4 ? , same host, same location, same elevation, 25- 1-1967, H. J. Egoscue (Egoscue); 1 ? , ex Per- omyscus manicidatus sonoriensis, Johnson Pass, 1797 m, 11-III-1969, H. J. Egoscue (Egoscue); 1 $ , same host, Wendover, 12- XII- 1964 (Egoscue); 1 ? , same host. Lake- side Mts., 16-III-1965 (Egoscue); 2 5,2 ? , Neotoma lepida lepida, Johnson Pass, 1872 m, 23-1-1969, H. J. Egoscue (Egoscue); 1 ? , same host. Little Granite Mt., 1418 m, 31-1- 1963, J. G. Bittmenn (Egoscue); 4 $ , same host, Wendover, 12-XII-1964 (Egoscue); 1 S , 1 ? , same host, Grassy Mt., 1590 m, 30-XI- 1967, H. J. Egoscue (Egoscue); 1 S , 2 ? , same host, W Stansbury Island, 1302 m. 29- 11-1968, H. J. Egoscue (Egoscue); 1 $ , ex Neotoma c. acraia, Johnson Pa.ss, 1830 m, 22- 1-1969, H. J. Egoscue (Egoscue); 1 (5,1 ? , ex Peromijscus crinitits pergracUis, N. Dug- way Mr., 1500 m, 27-11-1967, H. J. Egoscue (Egoscue); (Utah Co.) 1 ? , ex Neotoma cine- rea nest, Provo, 18-XI-1948, V. J. Tipson (BYU); 1 ? , same host, Buckley's Mine, Pro- vo, 24-XM949, D. M. Allied (BYU); 1 ^ , 3 ? , ex Neotoma cinerea (dung), Rock Canyon, Provo, 11-III-1949, V. J. Tipton (BYU); 1 $ , 3 $ , ex Neotoma cinerea, Provo Canyon, Provo, 12-XI-1949, H. Goldschmidt (BYU). Oregon: (Harney Co.) 6 S , 27 ? , ex Neo- toma cinerea alticola, 11.2 km S Crane, 1290 m. 25-XI-1968, H. J. Egoscue (Egoscue); 2 ? , ex Neotoma lepida nevadensis, 8 km S Crane, 1290 m. 25-XI-1968, H. J. Egoscue (Egoscue). Discussion.— Stenistomera alpina has a greater recorded distribution than the other two species of the genus and it probably oc- curs in all eleven western states. More than 90 percent of the specimens in our collection were associated with Neotoma lepida and most of them were collected during winter months. Although there is some slight varia- tion in the location of the spiniforms on the movable process of the clasper, most of our specimens show little variation in characters of taxonomic importance. Collection Data for Specimens Used for Illustrations CalU.stopsijIlus tciiuus campestri.s Holland: male and female, ex Peiomijsciis sp.. Alberta, Canada, 27-V-1949, G. P. Holland (Holland). CallistopsyUtis terintis deutenis Jordan: male and fe- male, ex Peromijscus tniei, 1.6 km S El Condor, Baja California, Mexico, 31-XII-1962, W. J. Wrenn (Traiib). CaUistopsyllus terinus tcrinus (Rothschild): male and female, ex Peromijsctis manicidatus, AEC-NRTS, Idaho, 22-III-1967 collectors unknown (BYU). ConorliinopsyUa nidicola Jellison: male and female paratypes, ex Neotoma sp. nest, Lawrence, Douglas Co., Kansas, ll-XI-1944, R. H. Beamer (Traub). ConorhinopsijUa stanfordi Stewart: male and female, ex Glaucomijs volans, Otsego Co., New York, 6-Xn- 1956, P. Connors (Benton). Megarthroglossus becki Tipton and Allred: male and female paratvpes, ex Neotoma cinerea nest, Buckley's Mine, Provo,' Utah Co.. Utah, 24X1-1949, D. M. Allred (BYU). Megarthroglossus bisetis Jordan and Rothschild: male and female, ex Neotoma micropus. Red Bluff Ranch, Chaves Co., New Mexico, 2,5-V-1967, B. Miller (Lewis). Megarthroglossus cavernicolus Mendez and Haas: male and female paratypes, ex Neotoma cinerea nest, cave W edge Valle Grande, Jemez Mts., Sandoval Co., New Mexico, 18-LX-1970, G. E. Haas et al. (Mendez). Megarthroglossus divisus (Baker): male and female, ex Seiurus fremonti nest, Elkhorn R. S., Wayne Co., Utah, 9-Vin-1952, M. Killpack et al. (BYU). Megarthroglossus divisus Wagner: male and female, ex packrat {Neotoma sp.) nest, 8 mi Creek, Ravalli Co., Montana, 29-XIM962, C. M. Senger (Senger). Megarthroglossus jamesoni Smit: male and female, ex Neotoma cinerea nest, Pine Nut Mts., Douglas Co.. Ne- vada, 25-in-195L E. W. Jameson (Jameson). Megarthroglossus procus procus Jordan and Rothsch- ild: male and female, ex packrat (Neotoma sp.) nest. Gla- cier, Whatcom Co., Washington, .\-1963, C. NL Senger (Senger). Megarthroglossus procus procus Jordan and Rothsch- ild: male ex "chipmimk, " Washoe Co., Nevada, X-1936, C. R. Eskey; female ex "chipmunk." Eldorado, Califor- nia, X-1936, C. R. Eskey. Megarthroglossus procus muiri .\ugustson: male holo- tvpe, ex Tamiasciurus d. albolimbatus, Tully's Hole, Fresno Co., California, 25-Vin-194L G. F. .4ugustson (.■\ugustson); female allotype, ex Sorex (Neo.) navigator. Tully's Hole, Fresno Co.,' California, 24-VHM941. G. F. .•\ugustson (.\ugustson ). Megarthroglossus sicamus Jordan and Rothschild: male paralectotype, ex Canis latrans. Eagle River, Si- canious, British Columbia, Canada, 6-L\-1903, G. F. Dippie (Smit); female ex Neotoma sp.. Pavilion Lake, British Columbia. Canada, 5-Vn-1950, K. B. (Holland). Megarthroglossus sierrae ,\ugustson: male holotype, ex Ochotona s. muiri. Cascade Valley, Fresno Co., Califor- nia, 27-VIII-194L G. F. Augustson (Augustson); female allotype, ex Tamiasciurus d. albolimbatus. Mammoth Lakes, Mono Co., California, 3LVII-19.39, G. F. August- .son (Augustson). December 1979 Tipton et al.: Anom lOPSYLLINAE (11 Figs. 103-108. Males, sternum IX; 103, Stenistomera alpina ; 104, S. macrodartifla; 105, .S. huhhardi, 106-108. claspers; 106, S. alpinci; 107, S macrodactyla; 108, S. hubbardi. 412 Great Basin Naturalist Vol. 39, No. 4 Megarthroglossus spenceri Wagner: male holotype {Megarthroglosstis pygmaeus), ex Neotoma cinerea occi- dentalis, Nicola, British Columbia, Canada, 25-VIII- 1932, collector unknown (Holland); female holotype {Megarthroglosstis spenceri), ex Oclwtona princeps, Ni- cola, British Columbia, Canada, 26-VIIM932, collector unknown (Holland). Megarthroglossus smiti Mendez: male and female paratypes, ex Neotoma lepida, Lynndyl, Millard Co., Utah, 17-XI-1951, Barnum, Moore, Melander and Clo- ward (BYU). Megarthroglossus tveaveri Eads and Campos: male and female paratypes, ex Neotoma mexicana. Weaver Ranch, Larimer Co., Colorado, 25-XI-1974 and 3()-X-1973 (re- spectively), E. G. Campos (Eads). Megarthroglossus wilsoni Mendez and Haas: male paratype, ex Eutamias minhntis nest, 0.8 km SE Red River Pass, Colfax Co., New Mexico, 28-Vin-1971, Haas and Wilson (Mendez); female paratvpe, ex Peromyscus mankulatus, 7.2 km NW Ft. Collins, Larimer Co., Colo- rado, 29-X-1969, collector unknown (Eads). Figs. 109-110. Males, aedeagus; 109, Stenistomcra hubbardi- 110, .S. manodactyla. December 1979 Tipton et al.: Anomiopsyllinae 413 Figs. 111-114. Males, aedeagiis; 111, Steni.stomera alpina. sternum VIII; 112, S. alpina, 113, S. macrodactyla; 114, S. hubbardi. 414 Great Basin Naturalist Vol. 39, No. 4 WESTERN NORTH AMERICA ^: DISTRIBUTION OF THE GENUS STENISTOMERA •S. alpina " S. macrodactyla * S. hubbardi Fig. 115. Distribution of Stenistomcm alpina, S. huhhardi and S. mdcwdacti/la. December 1979 Tipton et al.: Anomioi'syi.linae 415 StcttistoDicni (ilpiiui (Baker), male and female, ex Neo- toiiiti Icpidii, Mcrcmy, AEC-NHTS, Nye Countv, Ne- vada. 1 l-III-19(i(), collector unknown (BYU). Stenistoinera huhhurdi Egoscue, male and female, ex Pewmysctts inaniculdtus, 8.0 km S at Crane, Harnev County, Nevada, 23-X-19(S6, H.J. Ego.scuc. StcnistoDicra mdcwdacliild Cood, male and female, ex rcn>nujscu.s )n(iniciil(itiis. AEC>-NKTS, Bingham Countv. Idaho. 19X1-1967, collector unknown (BYU). Literature Cited .\LLREn. D. M. 1952. Plague important flea.s and mam- mals in Utah and the western United States. Great Basin Nat. 12(l-4):67-75. 1968. Fleas of the National Reactor Testing Sta- tion. Great Basin Nat. 28(2):78-87. .\mi.n, O. M. 1973. A preliminary survey of vertebrate ectoparasites in southeastern Wisconsin. J. Med. Entomol. 10(1):11()-111. AuGusTsoN, G. F. 1941. Contributions from the Los .An- geles Musemii Channel Islands Biological Survey. Three new fleas (Siphonaptera). Bull. S. Calif. Acad. Sci. 40(2): 101-107. 1942a. Ectoparasite-host records from the Sierran Region of east-central California. Bull. S. Calif. .\cad. Sci. 40(.3): 147-157. 1942b. i\ new flea from the Mojave Desert (Cali- fornia). Allan Hancock Found. Univ. S. Calif, pp. 138-146. 1943. Preliminary records and discussion of some species of Siphonaptera from the Pacific South- west. Bull. S. Calif. Acad. Sci. 42(2):69-96. 1953. Some new fleas (Siphonaptera) of western United States. Bull. S. Calif. Acad. Sci. 52(3): 119-126. 1955. Records of fleas from the Pacific southwest. Bull. S. Calif. Acad. Sci. 54(l):36-39. Baker, C. F. 1895. Preliminary studies in Siphonaptera. IV. Canadian entomol. 27(5): 130-132. 1898. Notes on Siphonaptera, with descriptions of four new species. J. New York Entomol. Soc. 6:53-56. 1904. A revision of American Siphonaptera, or fleas, together with a complete list and bibliogra- phy of the group. Proc. U.S. Natl. Mus. 27 (1361): 365-469, pis. 10-26. 1905. The classification of the American Siph- onaptera. Proc. U.S. Natl. Mus. 29(1417):121-170. Barnes, A. M., V. J. Tipton, and J. A. Wildie. 1977. 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Flea.s"of the state of Nevada. Bull. S. Calif. Acad. Sci. 48(.3): 115-128. Humphreys, J. G. 1967. Records of Ohio Fleas (Si- phonaptera). Ohio J. Sci. 67(3): 186- 190, loFF, I. G., AND O. I. ScALON. 1954. Handbook for the identification of the fleas of eastern Siberia, the Far East and adjacent regions. Medgiz, Moskva, Akad. Med. Nauk SSSR., 275 pp. Jackson, J. O., and G. R. DeFoliart. 1976. Relation- ships of the white-footed mouse, Perontysctis leucapus, and its associated fleas (Siphonaptera) in southwestern Wisconsin. J. Med. Entomol. 13(3): 351 -.356. Jameson, E. W., Jr. 1943. Notes on some fleas of New York State. Canadian Entomol. 75(9): 177. Jameson, E. W., Jr., and J. M. Brennan. 1957. An envi- ronmental analysis of some ectoparasites of small forest mammals in the Sierra Nevada, California. Ecol. Monogr. 27(l):45-54. Jellison, W. J. 1945. Siphonaptera: A new .species of December 1979 Tipton et al.: Anomiopsyllinae 417 ConorplunopsyUa from Kansas. J. Kaiis. Eiitomol. Soc. 18(3): 109-1 11. 1979. Personal coninuniication. Jellison, W. L., and L. CJlesnk. 1967. Index to tlie lit- erature of Siphonaptera of North America. Sup- plement 2, 1951-1960. U.S. Publ. Hlth. Serv., Natl. Inst. Hlth., Natl. Inst. Allergy and Infect. Dis., Rockv Mountain Lab., Hamilton. Montana. 406pp. Jellison, W. L., and N. E. Good. 1942. Inde.x to the lit- erature of Siphonaptera of North America. U.S. Publ. Hlth. Serv. Nat. Inst. Hlth. Bull. 178:1-193. Jellison, W. L., G. M. Kohls, .\nd H. B. Mills. 1943. Siphonaptera: species and host list of Montana fleas. Montana State Board of Entomol. Misc. Publ. 2:22. Jellison, W. L., B. Locker, and R. F. Bacon. 1953. In- dex to the literatiue of Siphonaptera of North America. Supplement 1, 1939-1950. U.S. Publ. Hlth. Serv. Natl. Inst. Hlth. Natl. Inst. Allergy and Infect. Dis., Rocky Mountain Lab., Ham- ilton, Montana. 246 pp. Jellison, W. L., .and C. M. Senger. 1973. Fleas of Mon- tana. Montana Agric. Exp. Sta., Montana State Univ., Bozeman, Montana, Res. Rept. 29:75 pp. 1976. Fleas of western North America except Montana in the Rocky Mountain Laboratory col- lection. Western Washington State College, Bel- lingham. Wash. p. 55-136. Jenkins, E., and A. W. Grundmann. 1973. The para- sitology of ground squirrels of western Utah. Hel- minthol. Soc. Washington 40(l):76-86. Jordan, K. 1933. A survey of the classification of the American species of CeratophyUns S. Lat. Novi- tates Zool. 39:70-79. 1937. On some North American Siphonaptera. Novitates Zool. 40(2):262-271. Jordan, K., and N. C. Rothschild. 1915. Contribution to our knowledge of American Siphonaptera. Ec- toparasites l(l):45-60. Layne, J. N. 1958. Records of fleas (Siphonaptera) from Illinois mammals. Nat. Hist. Misc. 162:1-7. Lewis, R. E. 1974. Notes on the geographic distribution and host preferences in the order Siphonaptera. Part 3. Hystrichopsyllidae. J. Med. Entomol. 11(2): 147-167. Mail, A., and G. P. Holland. 1939. Siphonaptera of western Canada in relation to sylvatic plague. Proc. 6th Pacific Sci. Congr. 5:125-128. Mendez, E. 1956. A revision of the genus Megarthro- ghssus Jordan and Rothschild, 1914 (Siphonap- tera: Hystrichopsyllidae). Univ. Calif. Publ. Ento- mol. 11(3): 159-192. Mendez, E., and G. E. Haas. 1972. A new flea of the genus Megartliroglosstts Jordan and Rothschild from New Mexico (Siphonaptera: Hystri- chop.syllidae: Anomiop.syllinae). J. Med. Entomol. 9(4):285-288. 1973. Megarthroglossm wilsoni, new species, with notes on the genus in New Mexico (Siphonaptera: Hystrichopsyllidae). Ann. Entomol. Soc. Amer. 66(5): 1129-1 139. Miller, B. E., D. L. Forcum, K. W. Weeks, J. R. Wheeler, and C. D. Rail. 1970. An evaluation of insecticides for flea control on wilil niainmals. J. Med. Entomol. 7(6):697-702. MoHLAN, H. B. 1954. Notes on the genus SUt^arlhw- glossiis (Siphonaptera, Hystrichopsyllidae) in San- ta Fe County, New Mexico, J. Parasitol. 40(4):446-448. 1955. Mammal fleas of Santa Fe County, New Mexico. Texas Rept. Biol. Med. 13(1 ):93- 125. Nelson, B. C, and C. R. Smith. 1976. Ecological effects of a plague epizootic on the activities of rodents inhabiting caves at Lava Beds National .Vion- uinent, California, J. Med. Entomol. 1.3(l):51-6i. OscooD, F. L., Jr. 1964. Fleas of Vermont. J. New York Entomol. Soc. 72(1 );29-33. Parker, D. D., and J. F. Howell. 1959. Host-flea rela- tionships in the Great Salt Lake Desert. J. Para- sitol. 45(6):507-604. Pollitzer, R. 1952. List of the wild rodent fleas which have been found plague infected in nature or proved to be susceptible to experimental in- fection. Bull. WHO 7:318-322, Annex 1. Poorbaugh, J. H., and H. T. Gier. 1961. Fleas (Si- phonaptera) of small mammals in Kansas. J. Kans. Entomol. Soc. .34(4): 198-204. Pratt, H. D., and H. E. Stark. 1973. Fleas of public health importance and their control. DHEW Publ. No. (CDC) 74-8267:42 p. Rail, C. D., D. L. Forcum, J. R. Wheeler, and E. E. Miller. 1969. Wild mammals and fleas of Red Bluff Ranch, New Mexico. J. Med. Entomol. 6(l):92-94. Rainey, D. G. 1956. Eastern woodrat, Xeotoina flori- dana: Life history and ecology. Univ. Kans. Publ. Mus. Nat. Hist. 9'(10):535-646'. Rothschild, N. C. 1903. A new British flea: Cerato- phijUus londiniensis. Entomol. Rec, pp. 64-65, 319-321. 1905. On North American Ceratophijllus. a genus of Siphonaptera. Nov. Zool. 12:153-174, pis. 4. 1915. Contribution to our knowledge of the Si- pho7iaptera fracticipita. Novitates Zool. 22: 302-308. Senger, CM. 1966. Notes of fleas (Siphonaptera) from Montana. J. Kans. Entomol. Soc-. 39(1): 105- 109. Simpson, G. G. 1964. Species density of North American recent mammals. Soc. Syst. Zool. 13(2):57-73. Smit, F. G. a. M. 1953. Descriptions of new and little- known Siphonaptera. Bull. Brit. Mus. (Nat. Hist.) Entomol. 3(5): 187-219. Spencer, C. J. 1936. A check list of the fleas of British Columbia, with a note on fleas in relation to saw- dust in homes. Proc. Entomol. Soc. British Co- lumbia 32:11-17. Stanford, J. S. 1931. \ preliminary study of Utah Si- phonaptera. Proc. Utah Acad. Sci. 8:1.53. 1944. More Utah Siphonaptera. Proc. Utah Acad. Sci., Arts, and Lett. 19-20:173-178. Stark, H. E. 1959. The Siphonaptera of I'tah. U.S. Dept. H.E.W., Public Hlth. Serv. 2.39 pp. 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. Ento- mol. 6(3):287-294. 418 Great Basin Naturalist Vol. 39, No. 4 Stewart, M. A. 1930. New Neartic Siphonaptera. Can. Entomol. 62(8): 175-180. Tipton, V. J. 1950. New distributional records for Utah Siphonaptera. Great Basin Nat. 10:1-4. Tipton, V. J., and D. M. Allred. 1951. New di,stribii- tion records of Utah Siphonaptera with the de- scription of a new species of Megarthroglonsus Jordan and Rothschild 1915. Great Basin Nat. 11:105-114. Tipton, V. J., and R. C. Saunders. 1971. A list of ar- thropods of medical importance which occur in Utah with a review of arthropod-borne disease endemic in the state. Brigham Young Univ. Sci. Bull, Biol. Ser. 15(2): 1-31. Tiraboschi, C. 1904. Les rats, les souris, et leurs para- sites cutanes dans leurs rapports, avec la propa- gation de la peste bubonic, .\rchiv. Parasit., (Paris), 8:161-349. Traub, R. 1953. Systematic notes on eyeless fleas. J. Washington Acad. Sci. 43:352-353. 1968. SmitcUa thambetosa n. gen. and n. sp., a re- markable "helmeted" flea from New Guinea (Siphonaptera, Pygiopsyllidae) with notes on con- vergent evolution. J. Med. Ent. .5(3):37.5-404. 1972. Notes on zoogeography, convergent evolu- tion and taxonomy of fleas (Siphonaptera), based on collections from Gunong Benom and else- where in southeast .\sia. II Convergent evolution. Bull. Brit. Mus. (Nat. Hist.) Zool. 2.3(10):.307-387, 20 plates. 1978. Personal communication. 1979. Personal commimication. Traub, R., and C. C. Hoff. 1951. Records and descrip- tions of fleas from New Mexico (Siphonaptera). Amer. Mus. Nov. 15.30:2.3pp. Traub, R., and V. J. Tipton, im]. jordauopsyllu allredi. a new genus and species of flea from Utah (Si- phonaptera). J. Washington .\cad. Sci. 41(8): 264-270. Wagner, J. 1889. Aphanipterologi.sche Studien. I. .Ana- tomic der vermipsvlla alaciut schimk. Soc. Ent. Rossica Horae 23:199-261. 1898. .\phanipterologische Studien, III. Ueber die Gattung Ptilcx imd Beschreibung neuer .\rten der Gattungen CeratophyUus, CtenopsyUa, Cera- topsylla. und Ti/phlopsyflo. Horae Soc. Ent. Ross. 3I:.555-.594, pis'. 8-10. ' Wagner, J. 19.30. Keys to the .\phaiiptera living on Mu- ridae. Mag. Parasitol. 1:97-192. 19.36. The fleas of British Columbia. Canadian Entomol. 68(9): 198-207, figs. l-IO. 1939. Klassen und Ordnungen des tierriechs. Fiinfter Bandi .\rthropoda, 3 Abteilung: Insecta XIII. Buch Teil f. Aphaniptera 1-114. 1940. Beitrag zur Kenntnis der Flohe von Bri.- Kolumbie. Zeitschr. Parasitenkund. 11(4): 463-468. Wehrle, L. p. 1953. A host index of Ai izoiia fleas. Pan- Pacific Entomol. 24(1):37-41. Whitaker, J. O., AND K. W. Corthum. 1967. Fleas of Vigo Countv, Indiana. Proc. Indiana Acad. Sci. 76:431-440.' Williams, L. A. and C. C. Hoff. 1951. Fleas from the upper Sonoran Zone near .\lbuquerque, .New Mexico. Proc. U.S. Nat. Mus. 101(3278):.305-313. Wise.man, J. S. 1955. The Siphonaptera (fleas) of Wyom- ing. Univ. Wvo. Publ. 19(1 & 2): 1-28. KARYOTYPES OF FOUR ARTEMISIA SPECIES: A. CARRUTHIl A. FIUFOLIA, A. FRICIDA, AND A. Sn\'ESCENS' E. Dui;l Arthur and C. l.ornizo Pope- .\bstract.- Artg»u,sia CMirtithii and A. frigida of the subgenus Artemisia and A. filifului and A. spincsccns of tht- subgenus Dracunculus all have chromosome numbers based on x = 9. Diploid (2n = 18) karyotypes of each species are composed of large, medium, and small chromosomes that are mainly metacentric and submetacentric. The individ- ual karyotypes are similar but distinctive. Artemisia filifolia's karyotype and chemistry is (juite similar to that of Section Tridentatae, but A. filifolia has significant morphological differences with respect to the Tridentatae. Arte- misia spinescens includes a tetraploid (2ri = 36) population as well as diploid populations. Karyofypic analysis of a tetraploid A. spinescens suggests that it is an autotetraploid, thus carrying out a common theme in Artcmi^i'i :.iii..- polyploidy). The genus Artemisia (Anthemideae, Com- positae) is principally a temperate northern hemisphere plant group (Good 1974, Bailey Hortorium Staff 1976). A few of its 250 spe- cies, however, extend to South America and southern Africa. Most Artemisia phylogenists have suggested an origin for Artemisia in Eu- rasia because of the preponderance of diverse species growing there and because most of its Anthemideae relatives occur there (Stebbins 1974, Cronquist 1978, McArthur and Plum- mer 1978, McArthur 1979). Beetle (1979) re- cently suggested an American origin for the genus. Even disallowing Beetle's hypothesis. North America is without question a center of diversity for Artemisia. Several Artemisia species complexes (Clausen 1951: groups of closely related plants capable of intragroup gene exchange) appear to be evolving in North America (Hall and Clements 1923, Keck 1946, Ward 1953, Beetle 1960, Estes 1969, McArthur and Plummer 1978). Artemisia has chromosome numbers based on x = 6,7,8, and 9 (Kawatani and Ohno 1964, Wiens and Richter 1966). By far the most common base number for Arternisia is x = 9, however, as it is for the whole of the Anthe- mideae (Persson 1974). Several Artemisia spe- cies and species complexes are composed of polyploid series. Euploid series based on x = 9 are most common (Table 1), but aneuploidy and amphiploidy based on other .t's are also known (Suzuka 1950, 1952, Kawatani and Ohno 1964). The euploid complexes may be flNfopolyploid with one basic genOnie or allo- polyploid with different genomes (Persson 1974) or somewhere in between— a segmental autopolyploid or allopolyploid (Stebbins 1971), in which case, genomes are partial!)' differentiated. Our work with Artetnisia has been mainly with the A. tridentata Nutt. complex ( = sec- tion Tridentatae) (Hanks et al. 1973, McAr- thur and Plummer 1978, McArthur et al. 1979, Welch and McArthur 1979). To better imderstand the Tridentatae, we have also looked at sympatric, non-Tridentatae, per- ennial Artemisia. This paper reports first publication of karyotypes of four non-Triden- tatae species. Two species represent the sub- genus Artemisia (A. carnithii Wood and A. frigicUi Willd.) and two represent the sub- genus Dracunculus (A. filifolia Torr. and A. spinescens D. C. Eaton), the third subgenus in Artemisia is Seriphidiuni, which is princi- pally Eurasian and North African. The sec- tion Tridentatae has been assigned to Seriphi- diuni, but the Tridentatae are probably independent of and parallel to the Seriphidia (McArthur and Plummer 1978). I official time and is therefore in the pubHc domain. Forest Service. U.S. Department of Agriculture, and Plant Breeder, Rice Researchers. Inc.. Glenn, California 95943. formerly 'Thisarticle was written and prepared bv U.S. Government emplo,^ .- ■ , . ,. <-, j lu \. uaam \^^,^ ^Research Genet..,, Intermountam Forest and Range Experiment Stat.on. Fores^t Service, y ^Depart-::',!^^ ^''^liri";^^^^^^ at the Inteniiountain Station's Shrub Sciences Laboratory, Prove, Utah I Biological Technician at the Shrub Sciences Laboratory. 419 420 Great Basin Naturalist Vol. 39, No. 4 Materials and Methods Table 2.— Artemisia accessions studied. Plant Materials.— The plant materials studied were from the native collection sites and from transplanted wildings at the Snow Field Station in Ephraim, Utah. Each collec- tion was assigned a culture number preceded by U to indicate its order of accession. Origi- nal locations of plant populations are given in Table 2. Voucher herbarium specimens for each accession have been deposited in the Shrub Sciences Laboratory Herbarium (SSLP). Karyotyping.— Seed was collected from open-pollinated plants at the Snow Field Sta- tion for A. carruthii, A. filifolia, and A. frig- ida, and from the natural populations for A. spinescens. Root tips from seedlings germi- Utah Taxon Culture collection site Elevation (ni) A. carruthii U4 Clear Creek Canyon, Sevier County 2,060 A. filifolia U7 Kanab, Kane County 1,510 A. frigida U9 Sunglow Park, Wayne County 2,070 A. spinescens U3 Gunnison, Sanpete County 1,550 A. spinescens U4 Ouray, Uintah County 1,420 Table 1.- - Euploid patterns in x = 9 Artemisia. Number of species- with 2n chromosomes Subgenus' 18 27 .36 54 72 18-36 18-45 18-54 18-72 18-90 36-54 References' 49 Keck (1946) Suzaka (1950, 1952) Arano (1962, 1963, 1968) Ehrendorfer(1964) Kawatani and Ohno (1964) Estes (1969) Korobkov (1972) Dracunculus Kawatani and Ohno (1964) Rousi (1968) Filatova(1971) Korobkov (1972) McArthur and Pope (1977) Seriphidium 15 1 Suzuka(1952) Kawatani and Ohno (1964) Filatova (1974a and 1974b) Persson (1974) Tridentatae' 73 45 21 Ward (1953) Taylor etal. (1964) McArthur and Plum nier (1978) McArthur (unpublished) 'See McArthur and Plummer (1978) and McArthur (1979) for historical development. The section Tridentatae has not been formally proposed as a sub- genus, but it is independent of and more or less parallel to the three recognized subgenera. 'The references and hence the number of species are not exhaustive, but are representative. December 1979 McArthur, Pope: Artemisia K AHYOTYI'ES 421 nated in petri dishes were pretreated with colchicine, fixed in 1:3 acetic alcohol, stained in acetocannine, and s(juashed on a micro- scope slide in Hover's solution (McArthur and Plummer 1978). Slides are stored at the Shrub Sciences Laboratory. Several seedlings of each accession were checked for chromosome numl)er (McArthiu- and Pope 1977), but karyotypes were pre- pared from one slide per accession-a slide with flat photogenic cells. Five randomly se- lected metaphase plates from each slide were photomicrographed (Fig. 1). Prints at a mag- nification of ;3120X were used to prepare the karyotypes in a manner slightly modified from that outlined in our earlier paper A * • « B* G .. D Fig. 1. Artemisia spp.: Photomicrographs of colchicine arrested metaphase plates of root tips (1400 X). .\, filifolia, culture U7, 2n = 18. B, fngiV/o, culture U9, 2n = 18. C, spinescens, culture U3, 2n=I8. D, spine.srem. culture U4, 2/1 = 36. The line in Fig. 1.4 defines the end of a ceil. 422 Great Basin Naturalist Vol. 39, No. 4 (McArthur and Pkimmer 1978). Each chromosome complement was assigned 100 arbitrary length units so that the actual mea- surement per chromosome was proportion- alized. For diploid accessions, large chromo- somes (L) were defined as those > 12.6 units, medium (M) as 9.6-12.6, and small (S) <9.6. For the tetraploid accession, these values were halved. The centromere position was recorded as the proportional length of the short chromosome arm with respect to the length of the long arm. Metacentric chromo- somes (M) were defined as those with a ratio >0.75, submetacentric (SM) as 0.50-0.75, and subteliocentric (ST) <0.50. Thus nine classes of chromosomes were possible: LST = large subteliocentric, LSM = large sub- metacentric, LM = large metacentric, MST = medium subteliocentric, MSM = medium submetacentric, MM = medium metacentric, SST = small subteliocentric, SSM= small submetacentric, SM = small metacentric. For preparation of the karyo- types of Table 3, chromosomes were grouped into the nine classes and paired by relative length and centromere position within each class. Interclass pairing was occasionally re- quired so that all chromosomes could be paired. In such cases, length was given prior- ity over centromere position. To confirm fit of pairing choices, each chromosome pair was visually inspected on the photo- micrographs. The five samples for each ac- cession were then averaged (x) for each chromosome pair and a standard error of the mean (se) computed. A means of measuring karyotype asym- metry (Wiens and Richter 1966), F percent is obtained for a karyotype by averaging the proportional length of each short chromo- some arm in respect to its long arm. In our case, F percent was obtained by halving the centromere position values of Table 3. As F percent decreases from a maximum of 50, a more asymmetric karyotype is indicated. Metaphase plates were available for A. car- mtJiii (Keck 1946) and A. frigida (Knaben 1968). We compared our F percent results Table 3. Karyotypic data for Artctuiski species. Taxon Chromosome Accession characteristics' 1 2 3 U4 L 12.75 ±.07 13.63 ±.24 12. 18 ±.10 C .75 ±.05 .90 ±.02 .93 ±.03 CC LSM LM MM U7 L 14.12±.28 13.75 ±.33 12.63 ±.14 c .71 ±.02 .92 ±.03 .92 ±.03 CC LSM LM LM U9 L 13.65 ±.33 10.59 ±.19 12.34 ±.22 C .92 ±.05 ..57 ±.07 .96 ±.01 CC LM MSM MM U3 L 13.40 ±.16 12. 11 ±.06 11.15±.20 C .78 ±.06 .71 ±.05 .64 ±.03 CC LM MSM MSM U4 L 6.71±.1I 6.30 ± .04 6.91 ±.11 C .81 ± .02 .82 ±.01 .63 ±.02 CC LM LM LSM L 10 11 12 U4 5.74 ±.03 5.62 ±.04 5.46 ±.06 C .89 ±.03 .85 ±.03 .84 ±.03 CC MM MM MM A. carruthii A. fill folia A. frigida A. spinescens A. spinescens A. spinescens 'Relative chromosome length (L), centromere position (C), and chromosome class (CC) December 1979 McArthur, Pope: Artemisia K VHYOTYI'FS 423 with values obtained from these previously published metaphase plates. In order to compare the diploid (2n = 18) and tetraploid (2m = 36) chromosome com- plements of A. spiiiescens, we used a paired t-test (Woolf 1968). The assiunption was that the relative lengths of the doubled pairs (1 with 2, 3 with 4, . . .17 with 18 of Table 3) would be significantly different (P<0.05) from tlie diploid pairs (1,2,. ..9) if the diploid chromosome complement had not doubled to form the tetraploids. We point out that be- cause size was the first pairing criterion, the possible doubled pairs' relative length of tet- raploids are systematically biased (e.g.,l>2, 3>4, ...17>18). For centromere position, the average of tetraploid pairs was compared to the diploids (1 and 2 with 1, 3 and 4 with 2, ...17 and 18 with 9). Results and Discussion The four diploid accessions have different, but not radically different, karyotypic pat- terns (Table 3, Fig. 1). Persson (1974:168) re- ported that the whole genus Artemisia has a relatively similar karyotype. We believe this tendency for a relatively similar karyotype is present in the genus, but some Artemisia taxa have quite different karyotypes (Filatova 1971, 1974a, 1974b), as well as probable aneuploid chromosome number reductions (Wiens and Richter 1966). Artemisia carruthii.- The accession of ,-\. carruthii that we examined has chromosome pairs as follows: 1 LSM, 1 LM, 5 MM, I SSM, and 1 SM (Table 3). Artemisia carruthii is a member of the A. hidoviciana Nutt. spe- cies complex of the subgenus .Vrtemisia. It occurs in inland western North America and is known only as a diploid (2n = 18). Keek's (1946) metaphase plate has an F percent of 37 as compared to our 44. Despite the appar- ent difference, the chromosomes are prob- ably similar. Estes's (1969) evidence (hybridi- zation along with meiotic chromosome pairing) supported autopolyploidy in the A. ludoviciami complex. Artemisia frigida.— Like A. carruthii, A. frigida is also a member of the subgenus Ar- temisia. Artemisia frigida, however, forms its Chromosome pair 4 5 6 7 8 9 F% 11.45 ±.15 10.96 ±.14 10.65 ±.15 10.43 ±.20 9.53 ±.29 8..39±.16 .90 ±.03 .99 ±.01 1.00±.00 .79 ±.05 .67 ±.06 .96 ±.02 44 MM MM MM MM SSM SM 9.99 ±.31 11.89 ±.28 10.95 ±.32 10. 10 ±.20 8..50±.38 9.05 ±.30 .50±.04 .90 ±.03 .84 ±.04 .97±.0I .38 ±.02 .86 ±.03 39 MSM MM MM MM SST SM 11.82±.16 .82 ±.04 11..32±.2I .98 ±.01 10.78 ±.18 .91 ± .08 10.53 ±.17 .96 ±.02 9..58±.13 .49 ±.02 9.36 ±.16 .85 ±.04 41 MM MM MM MM SST SM 10.62 ±.21 .73 ±.08 MSM 11.92 ±.19 .91 ± .05 MM 11.34±.19 .87 ±.03 MM 10.40 ±.13 .94 ±.05 MM 9.78 ±.20 .85 ±.09 MM 9.31 ±.14 .62 ±.07 SSM 39 6.36 ±.08 .62 ±.02 LSM 5.86 ±.12 .50 ±.05 MSM Chromosome 5.58 ±.13 .50 ±.05 MSM pair 5.41 ±.09 .60 ±.04 MSM 4.95 ±.06 .53 ±.03 MSM 6.0.3 ±.04 .91 ±.01 MM - 13 14 15 16 17 18 5.27 ±.10 .87 ±.03 MM 5.22 ±.10 .87 ±.03 MM 4.78 ±.06 .85 ±.05 SM 4.60 ±.03 .87 ±.02 SM 4.62 ±.10 .54 ±.06 SSM 4.58±.ll .41 ±.02 SST 36 424 Great Basin Naturalist Vol. 39, No. 4 own species complex. It ranges from Mexico through Alaska to Siberia (McArthur et al. 1979). Artemisia frigida is known only as a diploid (2n = 18) (Love and Love 1964, Kna- ben 1968, Kovanda 1972, Mulligan and Cody 1972, Hartman 1977, McArthur and Pope 1977). Its karyotype consists of 1 pair LM, 1 pair MSM, 5 pair MM, 1 pair SST, and a pair of SM chromosomes with an F percent of 41 (Table 3, Fig. lA). Knaben's (1968) cell had an F percent of 43. Artemisia filifolia.— This species is as- signed to the subgenus Dracunculus, but has no close relatives (Hall and Clements 1923). Beetle (1979) recently suggested a possible affinity between A. filifolia and the Triden- tatae. Our analyses show it to have pairs of chromosomes as follows: 1 LSM, 2 LM, 1 MSM, 3 MM, 1 SST, 1 SM with an F percent of 39 (Table 3, Fig. IB). This karyotype is quite similar to the Tridentatae— differing by two chromosome pairs. The mean Triden- tatae F percent is 38 (calculated from McAr- thur and Plummer 1978). When compared to the Tridentatae karyotype, A. filifolia has an extra LM pair in place of an MSM (McAr- thur and Plummer 1978). Kelsey and Shafiza- deh (1979) point out that the sesquiterpene lactone colartin is shared by A. filifolia and the Tridentatae. Our chromatographic data (Hanks et al. 1973 and unpublished data stored at the Shrub Sciences Laboratory) of phenolic compounds also show some sim- ilarities between the two taxa. Before any close relationship can be inferred, however, more definitive chemotaxonomic and system- atic study is required. The taxa differ widely in floral characteristics and wood anatomy (Moss 1940, McArthur 1979) and apparently do not hybridize despite areas of sympatric distribution. Artemisia spinescens.— This species is also currently assigned to the subgenus Dracim- culus (McArthur 1979). It differs from other species of Artemisia because it is both spring flowering and deciduous. Only four popu- lations have had chromosome numbers deter- mined (Powell et al. 1974, McArthur and Pope 1977). Of these, three were diploid (2n= 18) and one was tetraploid (2/i = 36). In the present study, the diploid karyotype has 1 LM, 3 MSM, 4 MM, and 1 SSM chromo- some pairs with an F percent of 39, whereas the tetraploid has a karyotype of 2 LSM, 2 LM, 4 MSM, 6 MM, 1 SST, 1 SSM, and 2 SM chromosome pairs with an F percent of 36. The tetraploid karyotype appears to be an approximate doubling of the diploid one. Our paired t-tests for relative length and centro- mere position showed tetraploid chromo- somes to be about what would be expected in doubling the diploid chromosomes. The ap- proximate doubling was especially true for relative length (P<0.50). The centromere positions were not significantly different from doubling (0.10>P>0.05), but did not indicate an exact doubling. Another measure of centromere position, F percent, was about the same for the diploid and tetraploid acces- sions (Table 3). Of particular interest were two pairs of LSM and SM and the single pairs of SST and SSM in the tetraploid. The LSM and SM could have been derived from the largest MSM and the smallest MM pairs (Pairs 3 and 8: Table 3) of the diploids by translocation. The SST and SSM pairs of the tetraploid could, with repatterning (e.g., pericentric in- versions), have been derived from the diploid SSM pair 9. Persson (1974) illustrated the analogous nonsimilar 4.v and 6.r polyploid grouping that occurs in the A. maritima L. complex. It is hard for us to visualize the tetraploid A. spinescens as anything other than auto- tetraploid. Artemisia spinescens has no close relatives. Although the diploid genome may have differentiated in various populations so that the tetraploid(s) may have arisen from hybrids between slightly different parents, the hypothetical differentiated diploids must surely have had a common soiuce in the re- cent past. Further support for the autotetra- ploid nature of tetraploid A. spinescens is the apparent tendency for autopolyploidy in the genus A)iemisia. Table 1 certainly suggests such a tendency. Furthermore both the A. lii- ckwiciana (Estes 1969) and the A. tridentata (Ward 1953, McArthur and Plummer 1978) complexes are riddled with autopolyploidy. Conclusions The four species examined in this report mirror much of the genus Artemisia's chromosomal picture. Their karyotypes are. December 1979 McArthur, Pope: Artemisia Karyotypes 425 in general, quite similar although there are distinctive differences. The four species are all .v = 9, as is most of Atiemisia. Artemisia spinescens shows apparent autopolyploidy, a phenomenon quite common in Artemisia. Ar- temisia filifolia has a karyotype quite similar to that found in the A. tridentata complex, but morphological and anatomical differ- ences do not support a close relationship be- tween these taxa. Acknowledgments This study was aided by federal fimds for wildlife habitat restoration through Pittman- Robertson W-82-R Project (Cooperators: In- termountain Forest and Range Experiment Station, USDA Forest Service, Ogden, Utah; and Utah State Division of Wildlife Re- sources, Salt Lake City, Utah). The Snow Field Station is cooperatively maintained by the above agencies, Utah State University, and Snow College. Literature Cited Arano, H. 1962. Cytological studies in subfamily Car- duoideae of Japanese Compositae VI. Karyotype analysis in the genus Artemisia. Bot. Mag. (Tokyo) 7.5:.356-368. 196.3. Cytological studies in subfamily Car- duoideae (Compositae) of Japan XIV. The karyo- type analysis on genus Artemisia (3). Bot. Mag. (Tokyo) 76:459-465. 1968. The karyotypes and geographical distribu- tion in some groups of subfamily Carduoideae (Compositae) of Japan. Kromosomo 72-73:2371-2.388. Bailey Hortorium Staff, L. H. 1976. Hortus third. Macmillan Publ. Co., Inc. New York. 1290 p. Beetle, A. A. 1960. A study of sagebrush, the section Trideutatae of Artemisia. Univ. Wyoming Agric. Expt.Sta. Bull. 368:1-83. 1979. Autecology of selected sagebrush species. In: G. F. Gifford, F. E. Busby, and J. K. Shaw (eds.). Sagebrush Ecosystem Symp., p. 23-26. Utah State Univ. Press, Logan. 251 p. Clausen, J. 1951. Stages in the evolution of plant spe- cies. Cornell Univ. Press. Ithaca, New York. 206 P- Chonquist, a. 1978. The biota of the Intermountain Re- gion in geohistorical context. Great Basin Nat. Mem. 2:3-15. Ehrendorfer, F. 1964. Notizen zur cytotaxonomic und evolution der gattung Artemisia. Osterr. Bot. Zeitschr. 111:84-142. EsTES, J. R. 1969. Evidence for autoploid evolution in the Artemisia hidoviciana complex of the Pacific Northwest. Brittonia 21:29-43. Fu.atova, N. S. 1971. De caryotipis sjH.'cieruin Arlemiaii e subgeiicre Dracunculus (Bess.) Rydb. in arenis vigenliuin. Bot. Mater. Gerb. Inst. Bot. Akad Nauk Ka/ahsk. SSR 7:46-49. 1974a. Ad stadio de caryotaxoMoiniciuii Artt'misii Kazachstaniae e subgenere Seriphidiiiin (Bess.) Rouy. Bot. Mater. Gerb. Inst. Bol. Akad Nauk Kazahsk. SSR 8:66-75. 1974b. Karyotaxonomy of two species of worm- wood of the subgenus Seriphidimu (Bess.) Rouy. Izvestiia Akademii Nauk Kazahsk. .SSR Sehia Bi- ologicheskaia 1 : 16-20. Good, R. 1974. The geographv oi flowering plants. 4th ed. Longmans Ciroup Ltd. London. .557 p. Hall, H. M., a.nd F. E. Cleme.nts. 192.3. The phyloge- netic method in taxonomy. The North American species of Artemisia, Cbrysothamnus and Atri- plex. Carnegie Inst. Wash. Publ. .326:l-.^55. IIanks, D. L., E. D. Mc.\rthi r, R. .Steve.ns, a.nd A. P. Pll'mmer. 1973. Chromatographic characteristics and phylogenetic relationships of Artemisia sec- tion Tridentatae. USD.\ For. Serv. Res. Pap. INT-14I. 24 p. Intermt. For. a«d Range Expl. Sta., Ogden, Utah. R\RTMAN, R. L. 1977. In: A. Love lOPB Chromosome number reports LVI. Taxon 26:257-274. Kaw.ata.m, T., and T. Ohno. 1964. Chromosome num- bers in Artemisia. Bull. Nat. Inst. Hyg. Sci. (Tokyo). 82:18.3-193. Keck, D. D. 1946. A revision of the Artemisia vulgaris complex in North .\merica. Proc. Calif. .\cad. Sci. 25:421-468. Kelsey, R. G., and F. Shafizadeh. 1979. Sesquiterpene lactones and systematics of the genus .\rtemisia (.\steraceae). P'hytochemistry 18: 1591-1611. Knaben, G. 1968. Chromosome numbers of flowering plants from central Alaska. Nytt Mag. Bot. 15:240-254. Korobkov, a. a. 1972. On the cytotaxonomical charac- teristics of some species of the genus Artemisia L. in the northeast of the USSR. Bot. Zurn. 57:1316-1.327. Kovanda, M. 1972. Somatic chromosome numbers for some Asteraceae. Rhodora 74:102-116. Love, .\., and D. Love. 1964. In: .\. Love and O. T. Sol- brig lOPB Chromosome number reports I. Taxon 13.99-110. McArthur, E. D. 1979. Sagebrush systematics and evo- lution. In: G. F. Gifford, F. E. Busby, and J. K. Shaw (eds.). Sagebrush Ecosystem Symp.. p. 14-22. Utah State Univ. Press, Logan. 251 p. Mc.\rthur, E. D., a. C. Blauer, A. P. Plimmer, and R. Stevens. 1979. Characteristics and hybridiza- tion of important intermountain shmbs 111. Sun- flower family. USDA For. Serv. Res. Pap. INT- 220. 82 p. Intermt. For. and Range Expt. Sta.. Ogden, Utah. McArthur, E. D., and A. P. Plummer. 1978. Biogeo- graphy and management of native western shrubs: a case study section Tridentatae of Arte- misia. Great Basin Nat. Mem. 2:229-243. McArthur, E. D., and C. L. Pope. 1977. In: A. L6ve lOPB Chromosome number reports LV. Taxon 26:107-109. 426 Great Basin Naturalist Vol. 39, No. 4 Moss, E. H. 1940. Interxylary cork in Artemisia with ref- erence to its taxonomic significance. Am. J. Bot. 27:762-768. Mulligan, G. A., and W. J. Cody. 1972. In: A. Love lOPB Chromosome number reports XXXV. Tax- on 21:161-166. Persson, K. 1974. Biosystematic studies in the Artemisia maritima complex in Europe. Opera Botanica .35:1-188. Powell, A. M., D. W. Kyhos, and P. H. Raven. 1974. Chromosome numbers in Compositae. X. Am. J. Bot. 61:909-913. Rousi, A. 1968. Cytogenetic comparison between two kinds of cultivated tarragon [Artemisia dractin- cidus). Hereditas 62:193-213. Stebbins, G. L. 1971. Chromosomal evolution in higher plants. Addison-Wesley Publ. Co., Reading, Mas- sachasetts. 216 p. 1974. Flowering plants: evolution above the spe- cies level. Harvard Univ. Press, Cambridge, Mas- sachusetts. 399 p. Suzuka, O. 1950. Chromosome counts in the genus Arte- misia. Jap. J. Genetics 25:17-18. 19.52. Chromosome numbers in Artemisia. Rep. Kihara Inst. Biol. Res. (Seiken Ziho) 5:68-77. Taylor, R. L., L. S. Marchand, and C. W. Crompton. 1964. Cytological observations on the Artemisia tridentota (Compositae) complex in British Co- lumbia. Can. J. Genet. Cytol. 6:42-45. Ward, G. H. 1953. Artemisia, .section Seriphidium, in North America, a cytotaxonomic study. Contr. Dudley Herb. 4:155-205. Welch, B. L., and E. D. McArthur. 1979. Feasibility of improving big sagebrush (Artemisia tridentata) for use on mule deer winter ranges. In: J. R. Goo- din and D. K. Northington (eds.). Arid land plant resources, p. 451-473. Texas Tech Univ. Press, Lubbock. 724 p. WiENS, D., AND J. A. RiCHTER. 1966. Artemisia patter- sonii: a 14-chromosome species of alpine sage. Am. J. Bot. 53:981-986. WooLF, C. M. 1968. Principles of biometry. D. Van Nos- trand Co., Inc., Princeton, New Jersey. 3.59 p. VARIATION IN LEAF ANATOMY AND CO^ ASSIMILATION IN SITANION HYSTRIX ECOTYPES' \Varren P. C:lai\ .\bstract.- Collections of Sitcmiun hijatiix known to differ in phenoiogical development. Iieinlit, dry matter pro- dnction, and total water use were examined for possible differences in leaf anatomy and in COo assimilation rates, Collections originating in warm, dry habitats produced the narrowest leaves with the fewest veins. Other anatomical characteristics examined were either not different among collections or the differences did not appear to he related to the original habitats. The CO2 assimilation rates were similar on per-unit weight basis; therefore total assimilation varied as a fvmction of plant size. Ecotypic variation within plant species has been studied by a number of investigators to better imderstand how plants adapt to their environment. The ecotype concept, or the genotypic response of a species to environ- mental factors, had its roots in the 19th-cen- tiiry work of Jordan (Quinn and Ward 1969). Some of the most notable advancements of knowledge were made by Turesson in the 1920s; by Clausen, Keck, and Hiesey in the 1940s; and by McMillan in the 1950s. Eco- typic responses to climate are often through variation in phenology and dry matter pro- duction, whereas responses to grazing are of- ten variation in growth forms and phenology. Responses to edaphic variation may be phys- iological rather than morphological or phe- nological. Within grass species, differences occur in numbers of vascular bundles, in stomate den- sity (Dobrenz et al. 1969a, 1969b), and in numbers of mesophyll cells (Wilson and Cooper 1969). Epidermal variations have been noted by Benson and Borrill (1969) and Gray et al. (1969). Photosynthetic and respi- ration rates have also been shown to vary within species (Klikoff 1968, Wilson and Cooper 1969). For grasses, however, it ap- pears to the author that most within-species variation in internal structure and photo- synthesis has been demonstrated among geno- types developed in plant-breeding programs, rather than among naturally occurring races. The differentiation in respon.se to climatic variation among Sitanion Injsthx ecological races occurs in phenological development, height, dry matter production, and respira- tory rate, but apparentlv not in water use ef- ficiency (Klikoff 1968; Clary 1975). The pur- pose of this studv of Sitanion hystrix leaf anatomy and CO2 assimilation rates was to determine whether differences in these char- acteristics appeared to be ecotypic responses to climate of the collection sites. Methods and Materi.^ls Clonal plant materials were collected in seven states to obtain samples from a wide variety of habitats and to provide ample op- portunity for ecotypic differences to occur among the populations studied. Collections were made in South Dakota, Nebraska, Colo- rado, New Mexico, Arizona, Utah, and Ne- vada (Fig. 1). Collection sites varied as nuich as 13°33' in longitude, 12° 14' in latitude, 1600 m in elevation, and 429 mm in annual precipitation (Table 1). These sites represent Merriam's ecological life zones of Upper Sonoran to Hudsonian (Lowe 1964). The plants were divided into ramets and studied in a transplant garden and a growth chamber. General procedures followed, as well as the study conditions, can be found in Clary (1975). Leaf Anato.my.— Leaf materials were gathered in the transplant garden during the first week of July for two years. Three leaves, This article wa. written and prepared by U.S. government employees on official time and U '^17'°'; '" '^;P"lJ|^h''°8;;;oT loc-..«l a. the ln.cTn„.nn...n Principal range scientist, USDA Forest Service, Intermountain Forest and Range Experiment Station, Ogden. Utah 84401. loca. ^Principal Station's Shrub Sciences Laboratory, Provo, Utah 84601 427 428 Great Basin Naturalist Vol. 39, No. 4 one from the center of each of three ran- domly selected plants, were obtained from individual collections. These were killed and fixed in Graf III solution. The leaf samples were sectioned and stained with a safranin- fast green schedule. The leaf cross sections were examined mi- croscopically. An ocular micrometer was used to measure three samples per leaf for: 1. Thickness of outer abaxial epidermal wall plus cuticle; 2. Total thickness of abaxial epidermal cells (Fig. 2); 3. Gross-sectional area (height times width) of substomatal cavity; 4. Gross-sectional area (height times width) of bulliform cells; and 5. Width of leaf cross section. The numbers of leaf veins were recorded in the following categories: (A) veins with a flattened top and a complete column of scle- renchyma fibers from upper to lower epi- dermis; (B) veins with a rounded top and an incomplete column of supporting tissue; and (G) veins with less than three-quarters the height of A and B category veins with an in- complete column of supporting tissue (Fig. 2). Thirty-five millimeter transparency pho- tographs were taken of the leaf cross sections. These were projected onto a dot grid and the proportion of structural tissue (vascular bun- dles and sclerenchyma fibers) was deter- mined. Silicone rubber impressions (Zelitch 1961) Fig. 1. Geographic distribution of the Sitanion hijatrix collection sites. December 1979 Clary: Leaf Variation 429 were made of the abaxial leaf surface of three plants in each collection in the transplant garden. The adaxial surface was not used be- cause deep venation obscured the stomata. A transparent positive was obtained by painting the nibber impressions with a thin film of cellulose acetate. Stomate density was deter- mined by counting the number per micro- scope field. An ocular micrometer was utiliz- ed to determine the average length and width of the stomatal apparatus (stoma, guard cells, and subsidiary cells). CO2 Assimilation.— Fifty-five potted ra- mets were used in a study of relative rates of CO2 assimilation. A movable plexiglass assim- ilation chamber was constructed that could accommodate 10 plants at a time. The cham- ber was set inside a plant growth room so imiform conditions could be provided. Tem- peratures inside the assimilation chamber were never lower than 21 C nor higher than 27 C. Each replication was exposed for two hours to 50 juC of Ci^Og. Four 2.54-cm leaf sections were immedi- ately taken from each ramet. Alcohol extrac- tions were made from two of the sections and beta particle emissions were counted with a CM tube and scaler. The remaining two sec- tions were oven dried and weighed. Beta counts per unit weight of leaf were then cal- culated. The counts were considered an ex- pression of CO2 as.similation activity. A second study of relative rates of (X)2 as- similation used Sitankm hy.sthx seedlings be- cause previous observations indicated that plants grown from seed may have more uni- form vigor than transplants. Seed was itsed from four sources: Blue Grade (.\rizona), Snow Bowl (Arizona), Long Park (Arizona), and McVey Burn (South Dakota) (Fig. 1). Seventy-two 1.9-liter cartons, 18 per source, were filled with a commercial potting mix and planted with eight seeds. Regular water- ing maintained a moist substrate. After two weeks, the plants were thinned to four per carton. Assimilation rates were 'measured at eight weeks. The procedure for extraction of O* from the seedlings was similar to that used for the ramets, except that the entire aerial portions of two plants from each carton were extract- ed and the aerial portions of the other two plants were oven dried and weighed. Table 1.— Description oi Sitanion hijstrix collection sites. Elevation Precipitation Climatic' Merriam Collection site Location (m) (mm) index life zone Snow Bowl (north central Ariz.) 111°42'W,35°20'N 2,980 739 0.93 HuHsonian Black Mesa (southwest Colo.) 107°29'W, 38°30'N 2,840 732 1.00 Hudsonian Bunker Hill (central Nev.) 117°07'W, .39°18'N 2,620 _ _ Upper Sonoran Turkey Plot (northwest N.M.) 108°33'W, 35°25'N 2,380 452 .49 Transition Long Park (central Ariz.) lir29'W, 34°53'N 2,260 574 .59 Transition Glorieta Mesa (north central N.M.) Ephraim (central Utah) 105°41'W, 35°22'N 111°36'W, 39°18'N 2,200 1,680 406 310 .43 .31 Upper Sonoran I'pper Sonoran McVey Burn (western S.D.) 103°34'W, 44°00'N 1,520 ,516 .62 Transition Fort Collins (northern Colo.) 1()4°54'W, 40°36'N 1,520 35.3 .37 Upper Sonoran Blue Grade (central Ariz.) 111°43'W, 34°4.3'N 1,440 396 .,30 Upper Sonoran Scottsbluff (western Neb.) 103°40'W, 42°00'N 1,380 356 .38 Upper Sonoran Santa Rita (southern Ariz.) 110°51'W,31°46'N 1,380 498 .30 Upper Sonoran ■Joint variable developed from relative growing season length and annual precipitation, and expressed as a proportion of the 430 Great Basin Naturalist Vol. 39, No. 4 Statistical Analyses.— Analyses of vari- ance were computed for leaf anatomy and as- similation data. A modified Tukey com- parison (Snedecor 1956) was used where appropriate to isolate significant differences among means. A climatic index was developed for the collection sites by multiplying an expression of growing season length times annual pre- cipitation, and expressing the result as a pro- portion of the maximum value. This posi- tioned each collection site on a scale of conditions from warm-dry to cool-wet. The numerical expression of growing season length was calculated by using the transplant garden as base 100. A value of one was added for each day later or subtracted for each day earlier than the transplant garden spring growth would normally be initiated at the collection sites (Hopkins 1918). This pro- cedure produced larger values for short growing seasons and smaller values for long growing seasons. Plant characteristics were studied in relation to the climatic index (Table 1). Results Leaf widths varied significantly among col- lections. The narrowest leaves occurred in those collections from warmer and drier sites. Collections of Sitanioii hystrix from such sites were previously reported to have lesser plant heights (Clary 1975), although the rankings for leaf width and maximum plant height are not highly correlated (r = 0.59). The relation- ship of leaf width to characteristics of the original collection sites has not been con- sistent among other grass species (Quinn 1969, Quinn and Ward 1969). The collections with the widest leaves gen- erally had the most veins per leaf. The total number of veins and vein categories A and C differed significantly among collections. The total veins per leaf appeared to be quite re- sponsive to conditions of the original collec- tion sites (Fig. 3). Plants from warm, dry hab- itats had the fewest veins per leaf. The numbers of veins were closely related to the climatic index of the collection sites and fol- lowed a consistent pattern in relation to Mer- riam's life zones. Collections with the highest number of category A veins (McVey Burn, South Da- kota; Long Park, Arizona; Turkey Plot, New Mexico; and Glorieta Mesa, New Mexico) were last to develop phenologically. These veins contain a greater concentration of structural tissue than do B and C category veins; thus, these results may support those of Fig. 2. Typical leaf cross section of Sitanion hi/strix: A = Category A vein; B = Category B vein; C = Category C vein; aec = abaxial epidermal cell; be = hulliforni cell; gc = guard cell; sc = subsidiary cell; and ssc = sub- stomatal cavity- December 1979 Clary: Leaf Variation 431 Christie and Mowat (1968). Those authors re- ported that the later the date of anthesis in Dactylis glomerata, the less digestible the plant to ruminant animals. Among collections, there were no differ- ences in vein density, thickness of outer abax- ial epidermal wall, thickness of abaxial epi- dermal cells, and cross-sectional areas of substomatal cavities and bulliform cells, or leaf thickness. Significant statistical inter- actions occurred between collections and time for several anatomy variables (epider- mal outer wall, leaf thickness, and B category veins), suggesting that the collections respond differently to differences in years. This may l)e to some extent the effect of timing growth to precipitation, although the differences in anatomy did not follow a predictable i)attern. Percent of supporting tissue, stomale den- sity, and stomate width differed significantly among the collections, but the differences ap- peared to have no geographical or ecological relationships. Other investigators have re- ported significant and often distinctive varia- tion in characteristics that had no apparent relation to environment (Ouiim 1969, Ouinn LIFE ZONES UPPER SONORAN TRANSITION HUDSONIAN UJ (WARM-DRY) (COOL-MOIST) CLIMATIC INDEX Fig. 3. Relationship of veins per leaf to two characterizations and a climatic index. of the original collection sites-Merriams life zones J2 Great Basin Naturalist Vol. 39, No. 4 ad Ward 1969). This may be the result of lo- ll genetic drift, particularly for character- tics that have little adaptive importance ad are not subject to strong selection pres- u-e. Carbon dioxide assimilation rates were leasured to obtain an index of photo- aithetic activity. Among collections, no dif- jrences were found in assimilation rates per nit weight of leaf in ramets or seedlings. As- milation of C'^ was very closely related to )tal weight of seedlings (r = 0.99). Assimilation rates were tested under only ae set of conditions. Respiration rates have een shown to differ among collections at )me temperatures, but not at others (Klikoff 968). The possibility thus exists that assimi- tion rates would have varied among collec- ons under different temperature regimens. Discussion Considerable disagreement exists on the alue of variations in leaf structure to CO2 jsimilation rates and efficiency of water use ^shton 1948, Maximov 1931, Shields 1950, ad Milthorpe 1961). The "xeromorphic" leaf )rm has often been considered important in lis regard. This form generally has a low irface to volume ratio, decreased cell size, licker cell walls, more compact network of eins, higher stomatal frequency, and thick- ning of the cuticle. The characteristics mea- ired in this study that most typify the xero- lorphic leaf form varied little among allections. In apparent harmony with this lack of sromorphic trends in arid habitat collections 'as the similarity among collections of assim- ation rates or of water requirements (Clary 975). The total CO2 assimilated and HgO anspired in a given period did not appear ) be a function of specific anatomical or hysiological adaptations, but largely a func- on of plant size, which varied among collec- ons. A major form of adaptation by Sitanion ystrix populations has been to match their ming of growth to the most favorable peri- d of the year, thus reducing the likelihood f severe moisture stress (Clary 1975). There- Dre, although the collections were from sites /ith widely differing climatic conditions. there may have been little selection pressure on most populations to differentiate a more xeromorphic leaf anatomy. Literature Cited AsHTON, T. 1948. Techniques of breeding for drought re- sistance in crops. Commonw. Bur. of Plant Breed, and Genet. Tech. Commun. No. 14. Benson, M., and M. Borrill. 1969. The significance of chnal variation in Dactijlis marina Borrill. New Phytol. 68:1159-1173. Christie, B. R., and D. N. Mowat. 1968. Variability of in vitro digestibility among clones of bromegrass and orchardgrass. Can. J. Plant Sci. 48:67-73. Clary, W. P. 1975. Ecotypic adaptation in Sitanion hys- trix. Ecology 56:1407-1415. DoBRENZ, A. K., L. N. Wright, A. B. Humphrey, M. A. Massengale, and W. R. Kneebone. 1969a. Sto- mate density and its relationship to water-use ef- ficiency of blue panicgrass (Panicwn antidotale Retz.).'Crop Sci. 9:354-361. DoBRENZ, A. K., L. N. Wright, M. A. Massengale, and W. R. Kneebone. 1969b. Water-use efficiency and its association with several characteristics of blue panicgrass (Panicum antidotale Retz.) clones. Crop Sci. 9:21.3-216. Gray, J. R., J. A. Quinn, and D. E. Fairbrothers. 1969. Leaf epidermis morphology in populations of Danthonia sericea complex. Bui. Torrey Bot. Club 96:.525-530. Hopkins, A. D. 1918. Periodical events and natural law as guides to agricultural research and practice. Mon. Weather Rev. Suppl. 9:.5-42. Klikoff, L. G. 1968. Temperature dependence of mito- chondrial oxidative rates of several plant species of the Sierra Nevada. Bot. Gaz. 129:227-230. Lowe, C. H. 1964. Arizona's natural environment. Uni- versity of Arizona Press, Tucson. 136 p. Ma.ximov, N. a. 19.31. The physiological significance of the xeromorphic structure of plants. J. Ecol. 19:27.3-282. Milthorpe, F. L. 1961. Plant factors involved in trans- piration. In: Plant-water relationships in arid and semi-arid conditions, Madrid Symp. Proc. Vol. XVI: 107-1 15. UNESCO, Paris. Qlunn, J. \. 1969. Variability among high plains popu- lations of Panicum virgatum. Bui. Torrey Bot. Club 96:20^1. Quinn, J. A., and R. T. Ward. 1969. Ecological differen- tiation in sand dropseed {Sporobohis cryp- tandrus). Ecol. Monogr. 39:61-78. Shields, L. M. 1950. Leaf xeromorphy as related to physiological and structural influences. Bot. Rev. 16:399-447. Snedecor, G. W. 1956. Statistical methods. Iowa State College Press, Ames. 5.34 p. Wilson, D., and J. P. Cooper. 1969. Effect of light in- tensity and CO2 on apparent photosynthesis and its relationship with leaf anatomy in genotypes of Lolium perenne L. New Phytol. 68:627-644. Zelitch, I. 1961. Biochemical control of stomatal open- ing in leaves. Proc. Natl. Acad. Sci. 47:1423-1433. INDEX TO VOLUME 39 The genera and species described as new to science in this volume appear in bold type in this index. Allred, Dorald M., and Arthur C. Cole, ar- ticle by, p. 97. Allred, Dorald M., and Vasco M. Tanner, ar- ticle by, p. 89. Annual energy budgets for three common ro- dent species in the northern Great Basin, p. 143. Anomiopsyllinae (Siphonaptera: Hys- trichopsyllidae), II. The genera Callistop- syUiis, ConorhinopsijUa, Megarthroglossus, and Stenistomera, p. 351. Ants from northern Arizona and southern Utah, p. 97. Armstrong, David M., article by, p. 199. Baumann, Richard W., article by, p. 241. Beetles from the environs of Lake Powell in southern Utah and northern Arizona, p. 89. Bee visitation of Phlox bryoides (Polemo- niaceae), p. 197. Bloss, Deborah Ann, and Jack D. Brotherson, article by, p. 161. Booth, Gary M., David A. Stewart, and Je- rold L. Petty, article by, p. 129. Boraginaceae of the southwestern United States, p. 293. Brothers, Donald R., article by, p. 195. Brotherson, Jack D., and Deborah Ann Bloss, article by, p. 161. Brotherson, Jack D., and Karen J. Brotherson, article by, p. 177. Brotherson, Jack D., Lorin E. Squires, Mark C. Whiting, and Samuel R. Rushforth, ar- ticle by, p. 245. Brotherson, Jack D., and Michael G. Skou- gard, article by, p. 44. Brotherson, Jack D., Shobha A. Jatkar, and Samuel R. Rushforth, article by, p. 15. Brotherson, J. D., and E. M. Christensen, ar- ticle by, p. 263. Brotherson, Karen J., and Jack D. Brotherson, article by, p. 177. Chemical composition of some important plants of southeastern Utah summer ranges related to mule deer reproduction, p. 122. Christensen, E. M., and J. D. Brotherson, ar- ticle by, p. 263. Clary, Warren P., article by, p. 427. Climates of fescue grasslands of mountains in the western United States, p. 284. Cnernonyx furvescens, p. 137. Cnemonyx protivorus, p. 137. Cnernonyx squamifer, p. 138. Cnemonyx vismiacolens, p. 138. Cole, Arthur C, and Dorald M. Allred, ar- ticle by, p. 97. Competition between harvester ants and ro- dents in the cold desert, p. 267. Competitive displacement as a factor in- fluencing phytoplankton distribution in Utah Lake, Utah, p. 245. Corthylus nanus, p. 138. Craig, T. H., and R. L. Gleason, article by, p. 274. Decreases of juniper woodland in the Utah and Salt Lake valleys since settlement, p. 263. Dendrocmnulns auctus, p. 139. Dendrocranuhis limbellus, p. 139. Dendrocrantdiis limitaris, p. 140. Dendrocranuhis modus, p. 140. Dendrocrantdiis pinguis, p. 140. Diatom floristics and succession in a peat bog near Lily Lake, Summit County, Utah, p. 15. Distribution of sculpins in the Clearwater River basin, Idaho, p. 59. Ecological and community relationships of Eriogonum corymbosum (Polygonaceae) in the Uinta Basin, Utah, p. 177. Ecological distribution of rodents in Canyon- lands National Park, Utah, p. 199. Emergence data and artificial rearing media for an aspen bark beetle, Trypophloeus popuU (Coleoptera: Scolytidae), p. 129. Feldhamer, George A., article by, p. 207. First record of Patapius spinosus in Idaho 433 434 Great Basin Naturalist Vol. 39, No. 4 and Nevada (Hemiptera: Leptopodidae), p. 195. Flinders, Jerran T., and Jeffrey S. Green, ar- ticle by, p. 88. Flora of the Lee Creek valley. Alberta, p. 277. Food habits of burrowing owls in south- eastern Idaho, p. 274. Gleason, R. L., and T. H. Craig, article bv, p. 274. Green, Jeffrey S., and Jerran T. Flinders, ar- ticle by, p. 88. Ground nesting and aggressive behavior of the Swainson's Hawk {Buteo swainsoni) p. 253. Harper, Kimball T., and John D. Shane, ar- ticle by, p. 219. Harper, K. T., and Jordan C. Pederson, ar- ticle by, p. 122. Heckmann, Richard A., Roger W. Mickelsen, and Rex C. Infanger, article by, p. 231. Higgins, Larry C, article by, p. 293. Homing by a pygmy rabbit, p. 88. Hylocurus clarki, p. 141. Hylocurus longipennis, p. 141. Infanger, Rex C, Roger W. Mickelsen, and Richard A. Heckmann, article by, p. 231. Influence of precipitation and temperature on ring, annual branch increment, and needle growth of White Fir and Douglas- fir in central Utah, p. 219. Jatkar, Shobha A., Samuel R. Rushforth, and Jack D. Brotherson, article by, p. 15. Jones, Stephen G., article by, p. 155. Jorgensen, Clive D., and Vichitra Mong- kolprasith, article by, p. 63. Jorgensen, Clive D., Dan S. Landeen, and H. Duane Smith, article by, p. 267. Karyotypes of four Artemisia species: A. car- rtithii, A. filifolia, A. frigida, and A. spin- escens, p. 419. Landeen, Dan S., Clive D. Jorgensen, and H. Duane Smith, article by, p. 267. McArthur, E. Durant, and C. Lorenzo Pope, article by, p. 419. McArthur, E. Durant, Charles F. Tiernan, and Bruce L. Welch, article by, p. 81. Maughan, O. Eugene, and Gary E. Saul, ar- ticle by, p. 59. Mickelsen, Roger W., Richard A. Heckmann, and Rex C. Infanger, article by, p. 231. Mongkolprasith, Vichitra, and Clive D. Jor- gensen, article by, p. 63. Mosher, James A., and Neil D. Woffinden, ar- ticle by, p. 253. Nash, Donald J., and David Wasserman, ar- ticle by, p. 192. Nearctic stonefly genera as indicators of eco- logical paramaters (Plecoptera: Insecta), p. 241. New synonymy and new species of American bark beetles (Coleoptera: Scolvtidae), Part VIII, p. 133. Peabody, Frederick J., article by, p. 1. Pederson, Jordan C, and K. T. Harper, ar- ticle by, p. 122. Petty, Jerold L., David A. Stewart, and Garv M. Booth, article by, p. 129. Phytoseiid predators of mite pests in Utah apple orchards, p. 63. Plilox longifolia Nutt. (Polemoniaceae) com- plex of North America, p. 1. Pope, C. Lorenzo, and E. Durant McArthur, article by, p. 419. Potential use of Great Salt Lake water for lobster culture, p. 231. Preliminary survey of raptor species in the Manti Division, Manti-La Sal National Forest, p. 155. Response of reptile populations to different land management practices on the Idaho National Engineering Laboratorv Site, p. 255. Review of tularemia in Utah and the Great Basin, p. 103. Reynolds, Timothy D., article by, p. 255. Rushforth, Samuel R., Lorin E. Squires, Mark C. Whiting, and Jack D. Brotherson, ar- ticle by, p. 245. Rushforth, Samuel R., Shobha A. Jatkar, and Jack D. Brotherson, article by, p. 15. Saul, Gary E., and O. Eugene Maughan, ar- ticle by, p. 59. Schreiber, R. Kent, article by, 143. Scaly todcs striatulus, p. 136. Shane, John D., and Kimball T. Harper, ar- ticle by, p. 219. Shaw, R. Keith, article by, p. 277. Sheldon, Andrew L., article by, p. 289. Skougard, Michael G., and Jack D. Broth- erson, article by, p. 44. Smith, H. Duane, Dan S. Landeen, and Clive D. Jorgensen, article by, p. 267. Squires, Lorin E., Mark C. Whiting, Jack D. Brotherson, and Samuel R. Rushforth, ar- ticle by, p. 245. December 1979 Index 435 Stark, Harold E., article by, p. 103. Stark, Harold E., Vernon J. Tipton, and John A. Wildie, article by, p. 351. Stewart, David A., Gary M. Booth, and Je- rold L. Petty, article by, p. 129. Stonefly (Plecoptera) records from the basin ranges of Nevada and Utah, p. 289. Studies in Nearctic desert sand dune Orthop- tera, Part XVI: A new black Stenopel- matiis from the Mescalero Sands, p. 226. Subspecies specificity of gall forms on Chrysotlianiniis nauseosus, p. 81. Tanner, Vasco M., and Dorald M. Allred, ar- ticle by, p. 89. Tepedino, V. J., article by, p. 197. Tiernan, Charles F., E. Durant McArthur, and Bruce L. Welch, article by, p. 81. Tinkham, Ernest R., article by, p. 226. Tipton, Vernon J., Harold E. Stark, and John A. Wildie, article by, p. 351. Variation in hemoglobin types in the Deer Mouse {Peromi/scus maniculatiis) along an altitudin..! gradient, p. 192. Variation in leaf anatomy and CO. assimila- tion in Sitanion hystrix ecotypes. p. 427. Vegetational response to three environmental gradients in the salt playa near (ioslien. Utah County, Utah, p. 44. Vegetation response to a moisture gradient on an ephemeral stream in central Ari- zona, p. 161. Vegetative and edaphic factors affecting abundance and distribution of small mam- mals in southeast Oregon, p. 207. Wasserman, David, and Donald J. Nash, ar- ticle by, p. 192. Weaver, T., article by, p. 284. Welch, Bruce L., E. Durant Mc.Xrthnr. and Charles F. Tiernan, article by, p. 81. Whiting, Mark C, Lorin E. Squires, Jack D. Brotherson, and Samuel R. Rushforth, ar- ticle by, p. 245. Wildie, John A., Vernon J. Tipton, and Har- old E. Stark, article by, p. 351. Woffinden, Neil D., and James A. .\losher. ar- ticle by, p. 253. Wood, Stephen L., article by, p. 133. NOTICE TO CONTRIBUTORS Original manuscripts in English pertaining to the biological natural history of western North America and intended for publication in the Great Basin Naturalist should be directed to Brigham Young University, Stephen L. Wood, Editor, Great Basin Naturalist, Provo. Utah 84602. Those intended for the Great Basin Naturalist Memoirs should be similarly directed, but these manuscripts are not encumbered by a geographical restriction. Manuscripts. Two copies of manuscripts are required. They should be typewritten, double spaced throughout on one side of the paper, with margins of at least one inch on all sides. Use a recent issue of either journal as a format, and the Council of Biology Editors Style Manual, Third Edition (AIBS 1972) in preparing the manuscript. 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Hayward, C. Cottam, A. M. Woodbury, H. H. Frost. $10. No. 2 Intermoimtain biogeography: A symposium. By K. T. Harper, J. L. Reveal, et al. $15. TABLE OF CONTENTS Boraginaceae of the southwestern United States. Larry C. Higgins 293 AnomiopsyUinae (Siphonaptera: HystrichopsylHdae), IL The genera CaUistopsylhis, Conorhinopsyllo, Megarthroglossus, and Stenistomera. Vernon J. Tipton, Harold E. Stark, and John A. Wildie 351 Karyotypes of four Artemisia species: A. cairuthii, A. filifolia, A. fiigido, and A. spin- escens. E. Durant McArthur and C. Lorenzo Pope 419 Variation in leaf anatoinv and Coj assimilation in Sitaniou Jn/strix ecotypes. Warren P. Clary '. ^ ' 427 Index to Volume 39 433 czncme , Bookbinding Co.. Inc. 100 Cambridge St. Charlestown. MA 02129 3 2044 072 231 012