IASELINE INVErciTORY OF MAMMALS, REPTILES, AND A I W H I B I A N S 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 m m a l s , 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 animals

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 1

Monument is indicated in Appendix I. The 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 <SO gr.) were

captured a total of 86 times, for a gross trapping success of

0.91 percent.

Table 3 details results of small mammal trapping. Alpha

diversity (species richness) across habitat-types ranged from one

to three. Mean number of individual small mammals of particular

species captured per transect ranged from 0 to 2.6. Total

individual small mammals captured per transect ranged from 0.25

on the prairie dog town to 4.2 in talus and cliff situations.

That converts to percentage trapping success of 0.12 to 2.1.

Mean biomass per transect ranged from only 6.6 in riparian

grassland to 75.0 in talus/cliff habitats.

This rate of capture is very low. For comparison, consider

trapping success in studies using comparable methods on a roughly

comparable fauna in roughly comparable habitats: Walker Ranch,

7000 it., Boulder Co., Colorado, 3.2 to 24..8% (Armstrong and

Freeman, 1984); Boulder Mountain Parks, Colorado, 4.5 to 22.0%

(Armstrong and Freeman, 1982); Main Elk Creek, ca. 6000 ft.,

Garfield Co., Colorado, 2 to 11% (Armstrong, unpubl.); Stagecoach

Reservoir site, Routt Co., Colorado, 5.0 to 13% (Armstrong, rC

unpubl. ) . Given the one-year duration of this project, reasons

~nderlying the 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, 2 3 ; wetland, aquatic, 18;

decidums woodland, 31; ponderosa pine woodland, 3 5 ; cliff-talus

shrub complex, 2 3 .

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 rodents 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.

I 1983. Mammals of the Northern Great Plains. Univ. Nebraska Press, Lincoln, xii + 379 pp.

Long, C. A. 1965. The m m a l s 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. - 17 December

4 February

5-8 June

4-29 August

1 September

1 December

12-14 December

16 December

1 February

1985 Proposal submitted

1986 Project funded

Preliminary field reconnaissance

Intensive field work

First Quarterly Report

Second Quarterly Report

Autumn Reconnaissance , Annual Report

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 .* Smooth Green Snake---is L,O =, \ \ut'pjllS Y

Wg ' 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* - ) - r u ' 1 - I /(

FAMILY SCIURIDAE--SQUIRRELS - fleast Chipmunk--Tamias minimus -ellow-bellied Marmot--Marmota flaviventris ~13-lined Ground 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 CRICETIDAE--NATIVE RATS AND MICE Western Harvest Mouse--Reithrodontomys megalotis mite-footed Mouse--Peromyscus 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 d e d Fox--Vulpes vulpes

FAMILY PROCYONIDAE--RACCOONS AND ALLIES L/~acc~on--~rocyon lotor

FAMILY MUSTELIDAE--WEASELS AND ALLIES dona-tailed Weasel--Mustela

/Striped skunk--Mephitis mephitis

L nl/ FAMILY FELIDAE--CATS d o b c a t - - A s rufus

FAMILY CERVIDAE--DEER tFdherican Elk--Cervus elaphus j

4 u l e 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 ~lpha diversity, and mean bGmass (grams) on transects --- in eahh of six habitat-types (number of replicate transects in parentheses). - - - 4,

SPECIES

HABITAT-TYPE

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 Wood Frog Spotted Frog Rubber Boa Montane Shrew Water Shrew Snowshoe Hare Pika

Frog

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 i n 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 const - Yellow-bellied Racer--Coluber rictor (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--Crotalus viridis, (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--Myotis evotis (Ch), P - Keen's ~yotis--Myotis keenii (B), 0, P - Small-footed Myotis--Myotis leibii (W), T, P - Little Brown at--Myotis lucifugus (w), 0, P - Fringed ~yotis--~yotis thysanodes (Ch), P - Long-legged ~yotis--Myotis volans (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-eared Bat--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--Lepus townsendii (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 Grasshopper Mouse--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 - Black-footed Ferret--Mustela niqripes (ext - Mink--Mustela vison (W), A - Badger--Taxidea taxus (W), S, M - Striped Skunk--Mephitis mephitis (W), S, 0 - Eastern Spotted Skunk--Spilogale putorius

, 0, PI M, A, T irpated)

FAMILY FELIDAE--CATS - Mountain Lion--Felis concolor (W), T I P - ~obcat--Felis rufus (W), P I T I 0

i-.22MILY CERVIDAE--DEER - ~ l k , or Wapiti--Cervus elaphus (W), M I P - Mule ~eer--0docoileus hemionus (W), P I 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)