IASELINE INVErciTORY OF , REPTILES, AND AIWHIBIANS OF DEVILS TOWER NATIONAL MONUMENT, WYOMING FINAL REPORT 15 June 1987

INTRODUCTION America's National Parks and Monuments provide u.nlque opportunities for biological research. The scenic beauty of many of the units of the National Park system is due to the mosaic of contrasting ecosystems they encompass; the beauty that delights 1 the casual visitor intrigues the ecologist. Because they are managed with careful attention to preservation of natural systems, many units provide the "baseline of normality" so important to understanding natural ecosystems. Finally, the interpretive programs of the Park Service provide an important

C- and effective forum for the dissemination of environmental knowledge. Public education is an important function of the academic ecologist and the National Parks provide a stimulating context for such outreach. Devils Tower National Monument comprises some 1346 acres in . Crook County, Wyoming. Despite its small size, the National Monument encompasses a wide array of ecological situations. The area is of particular interest to the ecologist and biogeographer because Devil's Tower and environs rise as an "islandn of montane znd saxicoline- habitat out of a "sea" of shortgrass prairie and sagebrush steppe. Studies of terrestrial mammals, reptiles, and amphibians in - northeastern Wyoming have been few. Baxter and Stone (1985) summarized knowledge of amphibians and reptiles in Wyoming, and Long (1965) compiled information on the mmals, including a nmher of records from Crook County. Turner (1974) reported on mammals of the Black Hills, including their western extent into Wyoming. Jones et al. (1983, 1985) summarized information on the natural history of the mammals of the Northern Great Plains. Accounts of all of mammalian species of Devil's Tower are included in those references. To my knowledge, the only studies of mammals in Devil's Tower National Monument are those reported by Halfpenny et al. (1977) and Owings and Owings (1977). The intent of this study was to investigate the ecological distribution of the terrestrial vertebrate fauna of Devils Tower National Monument. The functions of the study were (1) to aid in the interpretive program of the National Park Service; (2) to aid I in the resource management mission of the National Park Service; and (3) to provide the natural history baseline data upon which further ecological or evolutionary studies of the fauna of Devils Tower National Monument must be based. Surveys of ecological distribution of species are of practical importance and scientific interest. Figure 1 is a sketch of some potential relationships of ecological faunal surveys to other studies of basic and applied ecology. In order to protect and manage the environment of any area we must understand how the system operates. The first step toward such understanding is an inventory of organisms living there and some preliminary ideas of the ways they interact with their surroundings. METHODS Table 1 is a skeletal chronology of the project. Intensive field work was concentrated in August 1986, but at least some ~5 field observations were made in all seasons of the year and under a wide variety of field conditions. A total of 48 person-days were spent in the field. Studies were conducted with survey methods employed successfully in previous studies in Colorado (Armstrong, 1977s and Utah (Armstrong, 1979), and in use in on-going work in Rocky Mountain National Park (Armstrong, 1984). All methods were 1 designed to be as non-manipulative as possible. The only that must be captured for identification and'documentation are small mammals (< ca. 50 gr.). To study small mammals, a pair of transects (each with 25 stations, the stations set 5 m. apart and the transects spaced 10 m. apart) was established at each field .4 site. Each station was set with a Sherman livetrap. Traps were provided with cotton bedding and baited with rolled oats. Transects were run for four days and nights without pre-baiting. Interval of inspection depended on weather conditions, and was established to eliminate trap mortality. A total of 47 such transects was run, totalling 9400 trap- nights (47 transects x 50 traps x 4 nights). Small mammals captured were identified to species, sexed, weighed, aged, their reproductive condition and microhabitat noted, individually marked (by toe-clipping) and released. Numbers of animals per trapnight per species provides an index of relative abundance of species. Reptiles, amphibians, and larger mammals (deer, lagomorphs, - carnivores) were observed on plotless cruises through all major - habitat-types in all seasons. An attempt was made to capture bats by mist-netting (2 x 10 m. nets on 3 m. poles) in all habitat-types utilized, whenever weather conditions permitted, total of over 30 net-nights. Furthermore, an attempt was made to identify concentrations of roosting bats. Transects were located and reconnaissance cruises were conducted in all major habitat-types utilized in the National Monument, including upland meadow, oak wgodland, ponderosa pine woodland, riparian grassland, shortgrass prairie (prairie dog town), and shrub complex on talus/cliffs. Transects were marked temporarily with plastic surveyor's tape; all tape was removed at the conclusion of livetrapping. In addition to the systematic transects and opportunistic observations described above, interested and knowledgable Park Service personnel were interviewed, and wildlife observation files at Monument Headquarters were reviewed and evaluated. All work was coordinated with appropriate Park Service ~ersonnel,and conducted as unobtrusively as possible. To facilitate ongoing interaction between Park Service personnel and survey personnel, a checklist of the terrestrial vertebrates of

the National Monument was prepared in June I986 and provided to the Monument for use by interpreters and visitors.

Acknow1edqments.--For assistance in the field I thank Rick A. Adams, Bonni Gilbert, John D. Armstrong, and Susan L. Jessup. Vera Ramirez and Mary French of the Natural Science Program assisted C in administration of project funds; Virginia Hart was the principal contact for the project in the Office of Contracts end

Grants. A variety of Park Service personnel contributed to the work of the Survey. Special thanks are due Dick Guilmette, Homer Robinson, and Bill Pierce. Finally, I thank Dr. Kenneth L. Diem, Director, National Park Service Research Center, University of Wyoming for his help throughout this study. RESULTS AND DISCUSSION Ecoloqical Distribution and Alpha Diversity.--The potential natural terrestrial vertebrate fauna of Devils Tower National Monument is indicated in Appendix I. The1 list has been revised slightly from that presented in earlier reports, reduced from 86 to 81 species. The reduction is made possible by field acquaintance with the range of habitats present in the National Monument. Of the 81 species in the potential natural fauna of Devils Tower National Monument, six have been extirpated from the area within historic times (although the pronghorn still occurs nearby). Table 2 indicates species observed in the course of the present survey, either as animals or their certain sign. The list includes 35 species. Thus, some 47 percent of the potential natural fauna of the National Monument actually was observed on the Monument in the course of this survey. Given the fact that the list in Appendix I is a liberal one, and given the fact that some of the species listed (e. g., some bats, black bears, mountain lions) would be expected in the Monument only on an occasional basis, documentation of nearly half of the potential natural fauna should be considered quite successful. Additional concerted work on bats might increase the percentage of the fauna documented rather quickly; documentation of other species

A probably will be slower and more opportunistic. Studies of Small Mammals.--Many species of small mammals can be identified only by inspection in hand. This demands livetrapping. Livetrapping effort totalled about 9400 trap- rights (50 traps x 4 nights x 47 transects). A total of 48 individual small mammals (mammals usually weighing

Reservoir site, Routt Co., Colorado, 5.0 to 13% (Armstrong, rC unpubl. ) . Given the one-year duration of this project, reasons ~nderlyingthe low trapping success can only be speculated. Populations of small mammals do vary from year to year and fluctuations of numbers of some groups (e. g., microtines) are well known. Given the condition of the habitat (with an abundance of vegetation in most habitats, and an abundance of plants persisting through the winter bearing seeds) it is eifficult to imagine that small mammals were food-limited. Futhermore, predator pressure would seem to be minimal; no owl pellets and almost no coyote scat was picked/ up in the Monument, despite a number of observers continually alert to it. In the absence of salient biological factors, one is led to consider the physical environment. Most small mammals are active in winter. In temperate areas they often depend upon a blanket -4 of snow as insulation from cold. When such an insulating blanket is lacking, survival may be poor. The winter of 1985-86 was cold and dry, especially early in the season. This could have had a severe negative impact on overwintering populations. Ur-fortunately, of course, in the absence of any data on populations of small mammals in the growing season of 1985, this hypothesis cannot be tested. Bioqeoqraphic is the study broad distributional patterns in the living world. Often, the insights of biography can tell us of the history of the biota and can inform the planning process. Areography is the study of the shapes of geographical ranges of species. Areal patterns are patterns of the shapes and sizes "r, of geographical ranges. Areographic patterns of mammals of the Plains States were analyzed by Armstrong et al. (1986), and their m discussion is useful in the present context. Appendix I indicates the areal distribution of terrestrial vertebrates of Devils Tower Naticral Monument. Broadly speaking, based not on ecology but on the shape of their distributional range, the terrestrial vertekrates of Devils Tower National Monument can be assigned to one of the following areographic faunal elements (percentage of the fauna in each element in parentheses): widespread (39.5), campestrian, or Great Plains (18.5), cordilleran, or Rocky Mountain (8.6), boreo-cordilleran (7.4), eastern (9.9), Nevadan, or Great Basin (4.9), and Chihuahuan species, species occurring together in northern Mexico, 11.1). Appendix I also indicates the ecological distribution of - species in six habitat-types. Annotations are based on field observations and on Armstrong et al. (1986), Jones et al. (1983, 1985), and Baxter and Stone (1985). The number of species (alpha diversity) per habitat-type is as follows:

shortgrass prairie, 27; meadow, 23; wetland, aquatic, 18;

decidums woodland, 31; ponderosa pine woodland, 35; cliff-talus

shrub complex, 23. The relationship between areal and ecological distribution is of interest. Campestrian and Chihuahuan.species mostly make up the fauna of the arid shortgrass prairie. Chihuahuan bats are typical of ponderosa pine woodland and the cliff-side ecosystem. Boreo-cordilleran and cordilleran species are found mostly in meadows and ponderosa pine woodland. Eastern species are typical A of deciduous woodland. Many widespread species occur in several habitat-types, but some species of wetlands are also widespread geographically, albeit highly restricted ecologically. The biogeography of the Campestrian species is not particularly interesting, only because they are adapted to grassland habitats and their habitat at Devils Tower is essentially continuous at the present time with such habitats in nearly all directions. Likewise, species of wetlands and streamside habitats (including deciduous riparian woodlands) have more or less continuous (albeit via dendritic and sometimes quite f narrow corridors) habitat along watercourses across the Great - Plains.

' It is the montane habitats of ponderosa pine woodland and montane meadows that provide the most interesting biogeographic insights. Devils Tower rises above the Great Plains and is part of an "island" of forested habitats in a "sea" of grassland. Islands--literal or figurative--are of particular biogeographic interest. The question immediately arises, how did the fauna get to the island? Can the species involved get from the "mainland" of similar habitats elsewhere at the present time? Or is the "sea" too wide and too inhospitable, in which case the species on the island must have gotten to the island in earlier times when the "sea" was narrower or even non-existent. There is a considerable amount of theory in island biogecgraphy which need concern us only peripherally. Suffice it to say that traditional island biogeography theory (e. g., MacArthur and Wilson, 1967) may be inapplicable in this case because it provides an equilibrium model that assumes continual colonization (at rates depending on such variables as island size and distance from the "mainland" source area) and extinction (at rates depending on such variables as island size and consequent ecological diversity). In the case of habitat islands like Devils Tower and environs, a non-equilibrium model of island biogeography may be more applicable. Opportunity for colonization by montane habitat specialists may be non-existent. Local extinction would proceed without any possibility for replacement colonization. Biogeographically speaking, Devils Power and the Bear Lodge Mountains are a peripheral part of the Black Hills. However, the range of habitats about Devils Tower is somewhat narrower than that in the Black Hills region generally. Therefore, the fauna is relatively depauperate. For example, on the Black Hills there are patches of spruce-fir timber, the habitat of the southern red-backed vole (Clethrionomys qapperi) and the northern flying squirrel (Glaucomys sabrinus). Those species would be expected rarely if ever in the ponderosa pine woodland of Devils Tower. Furthermore, because it is an island, the Black Hills Region as a whole has a depauperate fauna relative to some other mountain systems. By way of comparison, consider Table 4. There the amphibians, reptiles, and mammals of montane, subalpine, and alpine habitats of the Bighorn Mountains, the Yellowstone-Teton region, and the the Black Hills are compared. Table 5 is a matrix of faunal resemblance. Below the diagonal of the matrix is indicated the number of species in common. Above the diagonal is a simple Faunal Resemblance Factor, calculated as 2C / N + 1 N , where N and N are the numbers of species in the two faunas 2 1 2 under comparison. It is not at all surprising that the faunas of the Yellowstone-Teton region are more similar than to each other than either of those faunas is to that of the Black Hills. The range of nigh elevation habitats on the Black Hills is smaller than on the Birhorns or in northwestern Wyoming and--we presume-- - extinction has taken place on the Black Hills that cannot have been replaced by re-colonization from the mountainous "mainland1' of the Rocky Mountains or the boreal forest of the Upper Midwest 1 and southern Canada. The upland fauna of the Black Hills (and of Devils Tower) is a relict of Pleistocene distributions that were once nearly or quite continuous across what is now unsuitable prairie habitats. They reflect times in the past when continental glaciation in the 6 Upper Midwest and alpine glaciation in the Northern Rocky Mountains were accompanied by a suite of environmental conditions cooler and moister than at present, conditions that allowed a far broader distribution of what now are montane and subalpine hzbitats. As suggested above, the fauna-of Devils Tower is even less extensive than that of the Black Hills--Devils Tower is on the periphery of the island and is thus "twice depauperate," a "subset of a subset" relative to places of seemingly similar ecology in the Mountain West. Furthermore, the cliff-talus habitats of Devils Tower have none of the rich suite of Chihuahuan species characteristic of A such habitats to the south, along the Front Range of Colorado, for example. Armstrong (1972, 1977b and elsewhere) has enphasized the importance of these Chihuahuan species to the C alpha diversity of faunas in the Southern Rockies and on the Colorado Plateau. There is no indication that such species ever have occurred in any number north of the South Platte drainage; for example, they are not present in the rich faunal record at Little Box Elder Cave in the North Platte Watershed of southern Converse County, Wyoming (Indeck, 1987). In the absence of cliff-talus specialists (other than a few Cordilleran species: the bushy-tailed wooprat and the yellow- bellied marmot), the fauna of distinctive cliff-talus habitats at Devils Tower is composed of euryecious Cordilleran species like the least chipmunk and widespread species like the ubiquitous deer mouse.

c Just because the fauna of Devils Tower is depauperate

relative to ecologically analogous sites to the west and south does not mean that the fauna is uninteresting or unimportant, of course. The fauna enriches the experience of the visitor and it contributes to the integrity of the overall mosaic of the environment of the Monument. Clearly, one of the most memorable experiences of many visitors to Devils Tower National Monument is the opportunity to observe black-tailed prairie dogs at close range. SUMMARY Devils Tower National Monument was established to preserve a remarkable geologic phenomenon. Preservation of the porphyric volcanic neck as a public trust has meant preservation of about 2 C square miles of the surrounding rims, valleys, terraces, cliffs, floodplans, and a reach of the Belle Fourche River. The fauna of amphibians, reptiles, and mammals of DEvils Tower National Monument comprises over 80 species, about half of which were observed in the course of a one-year survey. The fauna occupies a range of habitats, and the persistence of a fauna this rich depends upon the persistence of a complex mosaic of habitats: shortgrass prairie, montane meadows, ponderosa pine woodland, deciduous woodland (dominated by oak), streamside meadows and wetlands, and the cliff-talus shrub habitat. 1 The fauna of Devils Tower is a peripheral subset of the species typical of the Black Hills Region of- South Dakota and adjacent Wyoming. Widespread species occur there along with many species of the Great Plains and the eastern deciduous forest, In addition, several species typical of the Rocky Mountains are isolated in the Black Hills (including Devils Tower and the Bear Lodge Mountains), an insular relect of the Pleistocene ecology of the Northern Great Plains. BIBLIOGRAPHY Andersen, K. W., and J. K. Jones, Jr. 1971. Mammals of northwestern South Dakota. Univ. Kansas Publ., Mus. Nat. Hist., 19: 361-393. Armstrong, D. M. 1972. Distribution of mammals in Colorado. Monogr., Univ. Kansas Mus. Nat. Hist., 3: i-x + 1-415. Armstrong, D. M. 1977a. Ecological distribution of small mammals of the upper Williams Fork Basin, Grand County, Colorado. Southwestern Nat., 22: 289-304. Armstrong, D. M. 1977b. Distributional patterns of mammals in Utah. Great Basin Nat., 37: 457-474. Armstrong, D. M. 1979. Ecoloqical distribution of mammals of canyonlands National Park,-~tah. Great Basin Nat . , 39: 199-205. - Armstrong, D. M. 1984. Effects of the Lawn Lake Flood on the local distribution of mammals: preliminary report. Processed report to Rocky Mountain National Park, Estes Park, 25 pp. P Armstrong, D. M., J. R. Choate, and J. K. Jones, Jr. 1986. Distributional patterns of mammals in the Plains States. Occas. Papers Mus., Texas Tech Univ., 105:l-27. Armstrong, D. M., and G. E. Freeman. 1982. Mammals of the Boulder Mountain Parks. Processed report, City of Boulder, Department of Parks and Recreation, 133 pp. ------. 1984. Mammals ofWalker Ranch. Processed report, Boulder County Parks and Open Space Department, 12 pp. + appendices. Baxter, G. T., and M. D. Stone. 1985. Amphibians and reptiles of Wyoming, 2nd edition. Wyoming Game and Fish Dept., Cheyenne, 137 pp. f Brown, J. H., and A. C. Gibson. 1983. Biogeography. C. V Mosby, St. Louis, xi + 643 pp.

Halfpenny, J. C., J. S. Beckman, C. E. Fuenzalida, and M. Johnson. 1977. Reconnaissance of the of Devils Tower National Monument, Wyoming [Abstract]. J. Colorado- Wyoming Acad. Sci., 9 (1): 42-43.

Hoffmann, R. S., D. L. Pattie, and J. F. Bell. 1969. The distribution of some mammals in Montana. 11. Bats. J. Mamm., 50: 737-741. Hoffmann, R. S., and P. L. Wright, and F. E. Newby. 1969. The distribution of some mammals in Montana. I. Mammals other than bats. J. Mamm., 50: 579-604. Indeck, J. 1987. Sediment analysis and mammalian faunal remains from Little Box Elder Cave, Wyoming. Unpubl. Ph. D. dissertation, Univ. Colorado, Boulder, ix + 179 pp.

Jones, J. K., Jr., R. P. Lampe, C. A. Spenrath, and T. H. Kunz. 1973. Notes on the distribution and natural history of bats in southeastern Montana. Occas. Papers Mus., Texas Tech Univ., 15: 1-12. Jones, J. K., Jr., D. M. Armstrong, and J. R. Choate. 1985. Guide to mammals of the Plains States. Univ. Nebraska ?ress, Lincoln, xvii + 371 pp. Jones, J. K., Jr., D. M. Armstrong, R. S. Hoffmann, and C. Jones. 1983. Mammals of the Northern Great Plains. Univ. Nebraska I Press, Lincoln, xii + 379 pp. Long, C. A. 1965. The mmals of Wyoming. Univ. Kansas Publ., Mus. Nat. Hist., 14: 493-758.

HacArthur, R. H., and E. 0. Wilson. 1967. The theory of island biogeography. Monogr. Pop. Biol., Princeton Univ., 1: xi + - 1-203.

Owinqs,-. D. H., and S. C. Owinqs. 1979. Snake-directed behavior by blackltailed prairie dogs ( Cynomys ludovicianus). 2. Tierpsychol., 49:35-54.

Turner, R. W. 1974. Mammals of the Black Hills of South Dakota and Wyoming. Misc. Publ., Univ. Kansas Mus. Nat. Hist., 60: 1-178. - Table 1. Project chronology. 1985 17 December Proposal submitted 1986 4 February Project funded

5-8 June Preliminary field reconnaissance 4-29 August Intensive field work 1 September First Quarterly Report 1 December Second Quarterly Report 12-14 December Autumn Reconnaissance , 16 December Annual Report

1 February Third Quarterly Report 24-25 March Winter Reconnaissance

I 24-26 May Spring Reconnaissance Table 2.--Amphibians, reptiles, and mammals, observedin Devils Tower National Monument, Crook County, ~yoming,l986-1987.

------CLASS AMPHIBIA--AMPHIBIANS FAMILY BUFONIDAE--TOADS jcreat Plains Toad--Bufo cognatus FAMILY RANIDAE--TRUE FROGS vLeopard Frog--Rana pipiens CLASS REPTILIA--REPTILES FAMILY COLUBRIDAE--COLUBRID SNAKES Wellow-bellied Racer--Coluber constrictor -Hognosed Snake--Heterodon nasicus Wg .* Smooth Green Snake---is L,O=, \ \ut'pjllS Y '5 v8ullsnake--Pituophis melanoleucus vPlains Garter Snake--Thannophis radix , FAMILY VIPERIDAE--PIT VIPERS /Prairie Rattlesnake--Crotalus viridis CLASS MAMMALIA--MAMMALS FAMILY VESPERTILIONIDAE--COMMON BATS '--Little Brown Bat--Myotis lucifugus wBig Brown Bat--Epteslcus fuscus FAMILY LEPORIDAE--RABEqc,TS AND HARES Cottontail--Sylvilaqus cf . audubonig* -) -ru '1 -I /( FAMILY SCIURIDAE--SQUIRRELS- fleast Chipmunk--Tamias minimus -ellow-bellied Marmot--Marmota flaviventris ~13-linedGround Squirrel--Spermophilus tridecemlineatus -s ludovicianus ,,Fox Squirrel--Sciurus niqer WPine Squirrel--Tamiasciurus hudsonicus FAMILY GEOMYIDAE--POCKET GOPHERS northern Pocket Gopher--Thomomys talpoides FAMILY CASTORIDAE--BEAVER /Beaver--castor canadensis FAMILY --NATIVE RATS AND MICE -- megalotis mite-footed Mouse-- leucopus -Deer Mouse--Peromyscus maniculatus /Bushy-tailed Woodrat--Neotoma cinerea /Prairie Vole--Microtus ochrogaster FAMILY ERETHIZONTIDAE--PORCUPINES Acrcupine--Erethizon dorsatum FAMILY CANIDAE--DOGS AND ALLIES - coyote--Canis latrans ded Fox--Vulpes vulpes FAMILY PROCYONIDAE--RACCOONS AND ALLIES L/~acc~on--~rocyonlotor FAMILY MUSTELIDAE--WEASELS AND ALLIES dona-tailed Weasel--Mustela /Striped skunk--Mephitis mephitis

L nl/ FAMILY FELIDAE--CATS dobcat--As rufus FAMILY CERVIDAE--DEER tFdherican Elk--Cervus elaphus j 4ule Deer--0docoileus hemionus white-tailed Deer--0docoileus virqinianus ------"Identification tentative; field identification not documented by specimen. Table 3. Mean number of individual small mammals taken ~lphadiversity, and mean bGmass (grams) on transects ---in eahh of six habitat-types (number of replicate transects in parentheses). - - - 4,

HABITAT-TYPE

SPECIES

Tamias minimus 0.11 Spermophilus 13-lineatus 0.00 Feromyscus leucopus 0.30 -P. maniculatus 1.00 Onychomys leucogaster 0.00 Microtus ochroqaster 0.00 Alpha Diversity 3 Mean Number of Individuals Captured 1.41 - 0.45 0.50 Mean Total Biomass (grams) 27.3 8.4 27.0 T3ble 4. Occurrence of species restricted montane, subalpine, ---znd/or alpine habitatsin-- the Black Hills, the Bighorn- blountains --and the Yellowstone/Tetcn Region, Wyoming.

Boreal Chorus Frog Wood Frog Spotted Frog Rubber Boa Montane Shrew Water Shrew Snowshoe Hare Pika Yellow-pine Chipmunk Least Chipmunk Uinta Chipmunk .r- Yellow-bellied Marmot Pine Squirrel Northern Flying Squirrel Bushy-tailed Woodrat Heather Vole Water Vole Southern Red-backed Vole Long-tailed Vole Meadow Vole F:ontane Vole Western Jumping Mouse Black Bear Marten Ermine Lynx Moose Table 5. Faunal resemblance matrix for species listed in Table 4.

BIGHORN MOUNTAINS

YELLOWSTONE/TETONS

BLACK HILLS APPENDIX I

'.. ANNOTATED CHECKLIST OF POTENTIAL 'NATURAL FAUNA OF AMPHIBIANS, REPTILES, AND MAMMALSOF-DEV71I;S TOWER NATIONAL MONUMENT, CROOK COUNTY, WYOMING

NOTE: The following checklist includes all species known or suspected to occur (or to have occurred within the past century) in Devils Tower National Monument. The list includes all species known from Crook, Campbell, and Weston counties, Wyoming, and adjacent counties in Montana, South Dakota, and Nebraska, which occur in habitats represented in the National Monument. Areal distribution (in the sense of Armstrong et al., 1986) is indicated by an initial in parentheses, 9 follows: W = Widespread, not identifiable with any discrete, areographic faunal element; C = Campestrian; B = Boreo-cordilleran; Co = Cordilleran; Ch = Chihuahuan; E = Eastern; N-=Nevadan. Ecological distribution is tabulated in six habitat types, indicated by initials as follows: S = shortgrass prairie; M = meadow; 0 = oak woodland; P = ponderosa pine woodland; T = talus/cliff shrub complex; A = aquatic/wetlands.

CLASS AMPHIBIA--AMPHIBIANS FAMILY PELOBATIDAE--SPADEFOOTS -Plains Spadefoot--Scaphiopus bombifrons (C), S FAMILY BUFONIDAE--TOADS -Great Plains Toad--Bufo cognatus (C), S, 0 -- Xoodhousels Toad--Bufo woodhousei (C), S, 0 FAMILY HYLIDAE--TREE FROGS -Boreal Chorus Frog--Pseudacris triseriata (B), A FAMILY RANIDAE--TRUE FROGS -Leopard Frog--Rana pipiens (W), A FAMILY AMBYSTOMATIDAE--SALAMANDERS -Tiger Salamander--Ambystoma tiqrinum (W), A, M, 0 CLASS REPTILIA--REPTILES FAMILY TRIONYCHIDAE--SOFT-SHELLED TURTLES -Western Spiny softshell--Trionyx spinifera (C), A FAMILY EMYIDIDAE--POND TURTLES -Western Painted Turtle--Chrysemys picta (W), A FA-MILY CHELYDRI DAE--SNAPPING TURTLES -Common Snapping Turtle--Chelydra serpentina (E), A FAMILY IGUANIDAE--1GUANID LIZARDS -Sagebrush Lizard--Sceloporus graciosus (N), T, P -.Eastern Short-horned Lizard--Phrynosoma douqlassi (C), S FAMILY COLUBRIDAE--COLUBRID SNAKES -Yellow-bellied Racer--Coluber constrictor (E), M, 0 -Hognosed Snake--Heterodon nasicus (W) S, 0 -Milk Snake--Lampropeltis trianqulum (E), M, P, 0, S -smooth Green snake--0pheodrys vernalis (B), M -Red-bellied Snake--Storeria occipitomaculata (E), 0 -Bullsnake--Pituophis melanoleucus (W), 0, P, M, T -Plains Garter Snake--Thannophis radix (C), M, A, 0 Wandering Garter Snake--Thamno

FAMILY VIPERIDAE--VIPERS -Prairie ~attlesnake--Crotalusviridis, (C)-, S, T, P CLASS MAMMALIA--MAMMALS FAMILY SORICIDAE--SHREWS -Masked Shrew--Sorex cinereus (W), M, A -Merriam's Shrew--Sorex merriami (N), S, P -Montane Shrew--Sorex monticolus (Co), M -Dwarf Shrew--Sorex nanus (Co), T FAMILY VESPERTILIONIDAE--COMMON BATS -Long-eared ~yotis--Myotisevotis (Ch), P -Keen's ~yotis--Myotiskeenii (B), 0, P -Small-footed Myotis--Myotis leibii (W), T, P -Little Brown at--Myotis lucifugus (w), 0, P -Fringed ~yotis--~yotisthysanodes (Ch), P -Long-legged ~yotis--Myotisvolans (N), P -Silver-haired Bat--Lasionycteris noctivagans (W), 0, P -Big Brown Bat--Eptesicus fuscus (W), 0, P -Hoary Bat--Lasiurus cinereus (W), 0, P -Townsend's ~ig-earedBat--Plecotus townsendii (Ch), T, P FAMILY LEPORIDAE--=BITS AND HARES -Desert Cottontail--Sylvilaqus audubonii JCh), S -Nuttall's Cottontail--Sylvilaqus nuttallii (Co), P -White-tailed ~ackrabbit--Lepustownsendii (C), S -Black-tailed Jackrabbit--Lepus californicus (Ch), S FAMILY SCIURIDAE--SQUIRRELS -Least Chipmunk--Tamias minimus (W), T, P -Yellow-bellied Marmot--Marmota flaviventris (Co), T, P, 0 -13-lined Ground squirrel--Spermophilus tridecemlineatus (C), S -Black-tailed Prairie Dog--Cynomys ludovicianus (C), S -Fox Squirrel--Sciurus niger (E), 0 -Pine Squirrel, or Chickaree--Tamiasciurus hudsonicus (B), P, 0 -Northern Flying Squirrel--Glaucomys sabrinus (B), P FAMILY GEOMYIDAE--POCKET GOPHERS -Northern Pocket Gopher--Thomomys talpoides (Co), M, P -Plains Pocket Gopher--Geomys bursarius (C), S, M FAMILY HETEROMYIDAE--POCKET MICE AND ALLIES -olive-backed Pocket Mouse--Peroqnathus fasciatus (C), S -Hispid Pocket Mouse--Peroqnathus hispidus (Ch), S -Ord's Kangaroo Rat--Dipodomys ordii (Ch), S FAMILY CASTORIDAE--BEAVER -Beaver--Castor canadensis (W), A FAMILY CRICETIDAE--NATIVE RATS AND MICE -Western Harvest Mouse--Reithrodontomys meqalotis (Ch), S, M -Plains Harvest Mouse--Reithrodontomys montanus (C), S -White-footed Mouse--Peromyscus leucopuq (E), 0 -Deer Mouse--Peromyscus maniculatus (W), S, P, T, 0, M -Northern --0nychomys leucoqaster (~h),S -Bushy-tailed Woodrat--Neotoma cinerea (Co),- T -Prairie Vole--Microtus ochroqaster (C), M -Muskrat--0ndatra zibethicus (W), A -Sagebrush Vole--Lemmiscus curtatus (N), S FAMILY ZAPODIDAE-- JUMPING MICE -Meadow Jumping Mouse--Zapus hudsonicus (B), M, A FAMILY ERETHIZONTIDAE--PORCUPINES -Porcupine--Erethizon dorsatum (W), PI 0, T FAMILY CANIDAE--DOGS AND ALLIES -Coyote--Canis latrans (W), S, M, 0, P, T -Gray Wolf--Canis lupus--extirpated -Swift Fox--Vulpes velox ( C) , S -Red Fox--Vulpes vulpes (W), A, 0 -- Gray Fox--Urocyon cinereoarqenteus (E), 0, T FAMILY PROCYONIDAE--F?ACCOONS AND ALLIES -Raccoon--Procyon lotor (W), 0, A FAMILY URSIDAE--BEARS -Black Bear--Ursus arnericanus (W), 0, P -Grizzly Bear--Ursus arctos--extirpated FAMILY MUSTELIDAE--WEASELS AND ALLIES -Long-tailed Weasel--Mustela frenata (W), S , 0, PI M, A, T -Black-footed Ferret--Mustela niqripes (extirpated) -Mink--Mustela vison (W), A -Badger--Taxidea taxus (W), S, M -Striped Skunk--Mephitis mephitis (W), S, 0 -Eastern Spotted Skunk--Spilogale putorius FAMILY FELIDAE--CATS -Mountain Lion--Felis concolor (W), TI P -~obcat--Felis rufus (W), PI TI 0 i-.22MILY CERVIDAE--DEER -~lk, or Wapiti--Cervus elaphus (W), MI P -Mule ~eer--0docoileushemionus (W), PI M -white-tailed Deer--0docoileus virqinianus (w), 0, M FAMILY ANTILOCAPRIDAE--PRONGHORN -Pronghorn--Antilocapra americana (extirpated) FANILY BOVIDAE--CATTLE AND ALLIES -Sison--Bison bison (extirpated) -Sighorn Sheep--0vis canadensis (extirpated)