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LIBRARY

OF THE

Museum of Comparative Zoology

JM 13 UNIVERSITY OF KANSAS miscellaneous MUSEUM OF NATURAL HISTORY ^^^^^^^™I^nNo. 60

MUS. COMP. ZOOU LIBRARY Mammals of the ^p^ ^ ^ ^^^^ Black Hills of harvard UNIVERSITY South Dakota and Wyoming

By Ronald W. Turner

UNIVERSITY OF KANSAS LAWRENCE 1974 April 3, 1974 UNIVERSITY OF KANSAS PUBLICATIONS MUSEUM OF NATURAL HISTORY

The University of Kansas Publications, Museum of Natural History, beginning with volume 1 in 1946, was discontinued with volume 20 in 1971. Shorter research papers formerly published in the above series are now published as Occasional Papers, Museum of Natural History. The Miscellaneous PubUcations, Museum of Natural History, began with number 1 in 1946. Longer research papers are pub- Ushed in that series. Monographs of the Museum of Natural History were initiated in 1970. All manuscripts are subjected to critical review by intra- and extramural speciahsts; final acceptance is at the discretion of the publications conamittee.

Institutional libraries interested in exchanging publications may obtain the Occa- sional Papers and Miscellaneous Publications by addressing the Exchange Librarian, University of Kansas Library, Lawrence, Kansas 66045. Individuals may purchase separate numbers of all series. Prices may be obtained upon request addressed to PubUcations Secretary, Museum of Natural History, University of Kansas, Law- rence, Kansas 66045. University of Kansas Museum of Natural History

Miscellaneous Publication No. 60

April 3, 1974

Mammals of the Black Hills of South Dakota and Wyoming

By

BoNALD W. Turner

World HeaJfJi Organization Scientist Bojolali Plague Lahoratonj Bojolali, Central Java Indonesia

A dissertation subinitted in partial fulfdlment of the requirements of the degree of Doctor of Philosophy, The University of Kansas, 1971.

University of Kansas Lawrence 1974 University of Kansas Publications, Museum of Natural History

Editor : Linda Trueb

Managing Editor: William E. Duellman

Miscellaneous Publication No. 60

pp. 1-178; 14 figures Published April 3, 1974

Museum of Natural History University of Kansas Lawrence, Kansas 66045 U.S.A.

Printed by University of Kansas Printing Service Lawrence, Kansas CONTENTS

INTRODUCTION ....- . 6

Acknowledgments 6

Historical Resume 8

Methods and Materials 11

ENVIRONMENT 13

Geography 13

Geology and Physiography 13 Climate 22

Soils 23

Hydrography 25

Vegetation 26

Pleistocene History 29

Influence of Man on the Environment 38

ACCOUNTS OF SPECIES 40

Order Insectivora 40

Sorex cinereiis haijdeni Baird 40

Order Chiroptera 43

Myotis keenii septentrionaUs (Trouessart) 43

Mijotis leibii ciliolahrum (Merriam) 43

Myotis hicifugus carissima Thomas 47

Myotis thysanocles pahasapensis Jones and Genoways 49

Myotis volans interior Miller 51

Lasionycterus noctivagans (Le Conte) 52

Eptesiciis fusctis paUidtis Young 53

Lasiurus horealis horealis (Miiller) 55

Lasiurus cinereiis cinereus (Palisot de Beauvois) 56

Plecotus toivnsemlii pallescens (Miller) 57

Order Lagomorpha 59

Sylvilagiis auduboni l)aileyi (Merriam) 59

Sylvilagus floridamis similis Nelson 60

Sytvilagus nuttaUii grangeri (J. A. Allen) 61

3 Lepiis totvnse7idii campanius Hollister 62 Order Rodentia 63

Eutamias minimus poUidiis (J. A. Allen) 63

Eutamias minimus silvaticus White 64

Marmota floviventris dacota (Merriam) 68

SpermopJiilus tridecemlineatus paJJidus (J. A. Allen) 71

Cynomijs hidovicianus ludovicianus (Ord) 74

Sciunis niger rufiventer E. Geoffroy St.-Hilaire 75

Tamiasciurus hudsonicus dakotensis (J. A. Allen) 76

Glavcomijs sahrinus hongsi (Rhoads) 81

Thomomijs ialpoides nebulosus V. Bailey 83

PerognatJuis fasciatus oJivaceogriseus Swenk 86

PerognatJius liispidis paradoxus Merriam 87 Dipodomys ordii luteohis (Goldman) 87 Dipodomys ordii terrosus HoflFmeister 88 90 Reithrodontomys megalotis dychei J. A. Allen

Feromyscus leucopus aridulus Osgood 92

Peromyscus maniculatus nehracensis (Coues) 96 Neotoma cinerea orolestes Merriam 103

Cleithrionomys gapperi brevicaudiis (Merriam) 105

Microtus longicaudus longicaudus (Merriam) 108

Microtus ochrogaster haydenii (Baird) 110

Microtus pennsylvanicus insperatus (J. A. Allen) 114

Ondatra zihethicus cinnamominus (Hollister) 118

Rattiis norvegicus (Berkenhout) 119

Mus musctdtis Linnaeus 119

Zapus hudsonicus campestris Preble 120

Erethizon dorsatum bruneri Swenk 122

Order Carnivora 123

Canis lotrans latrans Say 123

Canis lupus irremotus Goldman 124

4 Viilpes viilpes reiialis Merriam 126

Ursus (unericanu.s americanus Pallas 126

Ursus arctos horribilis Ord 127

Frocijon lofor Jiirtus Nelson and Goldman 128

Musteki erminea muricus (Bangs) 129

alleni 130 Mustela frenota ( Mcrriam )

Mustela nigipes (Audubon and Bachman) 130

Mustela vison letifera Hollister 131

Taxidea taxus taxus (Schreber) 132

Mephitis mephitis hudsonica Richardson 132

FeUs concolor hippolestes Merriam 133

Lijnx canadensis canadensis Kerr 134

Lynx rufescens pallescens Merriam 135

Order Artiodactyla 135 Cervns canadensis canadensis Erxleben 136

OdocoiJeus hemionus hemiomis (Rafinesque) 137

Odocoileus virginianus dacotensis Goldman and Kellogg 139

Antilocapra americana americana (Ord) 141

Bison bison bison (Linnaeus) 144

Oreamnos americanus missouJae J. A. Allen 145

Oris canadensis auduhoni Merriam 147 SPECIES OF UNVERIFIED OCCURRENCE 148

Species Incorrectly Reported from the Black Hills 148

Species of Uncertain Status in the Black Hills 149 FACTORS INFLUENCING DISTRIBUTION AND SPECIATION 153

Mammalian Distributional Patterns 153

Origin of the Recent Mammalian Fauna of the Black Hills 155

Speciation and Geographic Variation 162 SUMMARY 165 LITERATURE CITED 166 INTRODUCTION

The Black Hills have been described from outside North America. Also in- as a mountainous island surrounded by cluded in the accounts are 25 species in Black Hills is a sea of grass. The mesic climate, conif- whose occurrence the erous forests, rugged and dissected to- questionable or undocumented at pres- pography, and diversity of geological ent. structures and edaphic features in the Black Hills contrast sharply with the ACKNOWLEDGMENTS adjacent Northern Great Plains, which are characterized by semi-arid to arid Many persons have contributed di- grasslands and gently rolling topography. rectly or indirectly to the completion of In some areas, general zones of transition this paper. I am particularly grateful tend to compromise the distinctiveness of to the following persons for the loan of these two physiographic entities. In spite specimens, for information concerning of the northern and montane affinities of specimens, or for permission to examine the Black Hills, the indigenous mamma- specimens: S. Anderson, K. F. Koopman, lian fauna is heterogeneous in origin. A and R. G. \^an Gelder, American Mu- definitive biogeographical analysis of the seum of Natural History; J. C. Moore, mammalian fauna has not been at- Field Museum of Natural History; R. R. tempted prexiously. Mansfield, Minnilusa Historical Museum; The principal purposes of this report W. H. Burt and E. T. Hooper, Museum are: 1 to delimit and describe the mam- of of D. ) Zoology, University Michigan; J. malian fauna of the Black Hills of South Druecker and ]. S. Findley, Museum of Dakota and Wvoming (43 10'-44"50' N Southwestern Biology, University of lat.; 103 20'- 104 50' W long.) as a nat- New Mexico; R. A. Martin, Department ural zoogeographic unit; 2) to describe of Biology, South Dakota School of the autecology and distributional pat- Mines and Technology; D. W. Block, in terns of each mammalian species the E. J. Hugghins, and R. J. Walstrom, Hills; 3) to discuss the geographic varia- Department of Entomology and Zoology, tion and inferred speciation of these South Dakota State University; B. Har- mammals; and 4) to analyze and inter- rell. Department of Biology, Uni\'ersity pret the probable biogeographic affinities of South Dakota; C. O. Handley, Jr., of various species in light of proposed D. A. Schlitter, and H. W. Setzer,' U. S. changes in late Pleistocene and Holocene National Museum; R. H. Manville and

environments. Thus, this study repre- J. Paradiso, U. S. Fish and Wildlife Serv-

sents a of Collection ; C. A. synthesis systematic, zoogeo- ice ( Biological Surveys ) graphic, ccologic, and historic factors McLaughlin, Department of Zoology, and their bearing on the contemporary University of Wyoming; and N. R. Whit- mammalian fauna of the Black Hills. ney, Rapid City, South Dakota. I am The Recent Black Hills mammalian indebted to Park Superintendent W. D. fauna comprises 62 species in 44 genera Hotclikiss and Chief Park Naturalist and six orders. Three of the species have G. B. Robinson of Wind Cave National been extirpated by man and subse- Park for pro\iding physical facilities and quently reintroduced to the Black Hills access to records on file at the Park. I from other areas. Four other species wish to extend special recognition to Mich- (indicated by an asterisk in the Con- J. A. King, Department of Zoology, tents) are adventives; of these, two were igan State Uni\'ersity, for making a\ail- introduced from North America and two able his unpublished field notes and

6 TURNER: MAMMALS OF THE BLACK HILLS supplementary observations which great- pertaining to the larger fur-bearers in ly enhanced my study. I also wish to their respective districts. I am especially thank R. to Assistant Barbour and W. H. Daxis, De- grateful Chief Ranger J. F. partment of Zoology, Uni\ ersity of Ken- Devenport, Wind Cave National Park, tucky, for assistance in the field in the for contributing his firsthand knowledge summer of 1968. of wildlife within the Park and adjacent Some 46 persons associated with the regions. The advice of A. C. Meland, Soils and Water Conservation Museum of Natural History of The Uni- Service, the characteristics versity of Kansas participated in field concerning edaphic of work in the Black Hills from 1947 to the Black Hills region is appreciated. A number of in the U. S. Forest 1967. I want especially to recognize S. personnel Anderson, R. H. Baker, E. L. Cockrum, Service, Black Hills National Forest, ad- E. R. vanced in innumerable Hall, J. K. Jones, Jr., C. A. Long, my study ways. these I would mention and H. W. Setzer, whose expeditions Among especially resulted in specimens and comprehen- I. Case, W. D. Cloud, F. Fichtner, A. W. D. G. K. C. sive field notes used in preparation of Jones, Kocer, Roby, Scholz, and C. Windsor. I to all this report. J. am indebted those listed above and others unnamed, For identification of ectoparasites, I who of their time and am indebted to K. C. Emerson, U. S. gave generously efforts in behalf. Army; E. Garret, Bishop Museum, Hono- my The National lulu; C. E. Hopla, Department of Zool- Science Foundation ogy, University of Oklahoma; G. M. (through fellowships to student partici- for field work in Kohls, Rocky Mountain Laboratory, pants June and July Hamilton, Montana; R. B. Loomis, De- of 1965 and 1967 under Grant GE-7739), of Beach Cali- The Museum of Natural the partment Biology, Long ( History, of fornia) State College; B. McDaniel, De- Department Zoology, and the Com- mittee of and partment of Entomology and Zoology, Systematics Evolutionary Grant at South Dakota State University, Brook- Biology (NSF GB-4446X) The of a ings, South Dakota; B. V. Peterson, Can- University Kansas, and Watkins Mu- seum of Natural all ada Department of Agriculture, Ottawa; History Grant, R. D. Price, Department of Entomology, helped to defray cost of field operations. A the Fish, and Wildlife, University of Minne- computer grant from Department of and enhanced sota; the late R. L. Usinger, Department Systematics Ecology statistical of the of Entomology, University of California analysis data. (Berkeley); N. Wilson, Department of I am grateful to other friends and Biology, University of Northern Iowa, associates, especially Elmer C. Birney, Cedar Falls; and W J. Wrenn, Depart- John B. Bowles, Hugh H. Genoways, and ment of Entomology, The University of Carleton J. Phillips, who have given of Kansas. their time and criticisms, and I am The cooperation of the South Dakota obliged to Barry L. Siler for preparing the illustrations. Department of Game, Fish and Parks is To my wife, Barbara M. I for her fully appreciated. Scientific collecting Turner, give special recognition and permits were kindly issued by V. Johnson patience perse\'erance during the final of the and for and J. W. Sprague. Chief of the Division phases project typ- of the first draft of the Game Management, J. Popowski, and ing manuscript. Similar is to H. State Game Biologists, L. Petersen and recognition given Mary Michener for the final A. H. Richardson, contributed significant reading proof. information concerning the artiodactyls I am indebted to Robert S. Hoffmann and carnivores in the Black Hills region. and Ronald McGregor who supported State Game Wardens R. Butterfield, O. my investigation through encouragement Meadows, C. Webster, and E. L. Woods and contribution of crucial counsel. In also donated specimens and information addition, Wakefield Dort, Jr., and A. W. MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Kuchler read and criticised portions of Executive Documents of Congress. Ad- the manuscript. I extend my thanks to ditional information can be obtained

Howard J. Stains, Southern IlHnois Uni- from the collections and publications of versity, who first instilled in me an in- the South Dakota State Historical So- terest in zoology, and through his teach- ciety, Pierre, South Dakota 57501. and ing personal example gave great Francois and Louis-Joseph Veren- to formal education. impetus my drye, two French explorers, entered the Especially instrumental in the forma- Black Hills in early February of 1743; tion and completion of my study was J. they were probably the first white men Knox Jones, Jr., who gave continued to penetrate the region. Although Meri- guidance throughout the duration of the wether Lewis and William Clark did not project, devoted many arduous hours to enter the Hills proper, a French trader, counseling, assisted in field problems, Valle, encountered the explorers near and aided in preparation of the com- the mouth of the Cheyenne River and pleted manuscript. His unselfish assist- told them of the "Black Mountains" to ance is acknowledged with sincere grati- the west. Having received similar re- tude. Professor Jones pro\'cd himself to ports while camped at the mouth of the be capable of considerable patience, the Bad River in 1833, Prince Maximillian depths of which only I can truly appreci- termed this pine-clad range the "Black ate. Hills" (Thwaites, 1906). He also in- cluded the Killdeer Mountains and Little HISTORICAL resume: Missouri River Badlands of North Da- kota this under designation ( Bailey, Little is known about prehistoric man 1927:25). Both Hunt's American Fur in the Black Hills area. Archeological Company Astoria Expedition (1811) and findings from several sites indicate that Jedediah Smith's expedition of 15 fur early inhabitants of the area (7000-9000 traders (1823), passed through parts of BP) were bison hunters organized so- the Black Hills. Additionally, there was cially into small hunting bands, each a number of other traders and trappers composed of several cooperating families who traxersed the Hills during the first (Black Hills Area Resource Study, 1967: half of the nineteenth century, but these 21). Several Indian tribes (in chrono- men left little record of their passing. logical sequence: the Poncas, Kiowas, Fur trading posts were established near Crow, and Cheyennes) lived near the the mouth of the Belle Fourche Ri\'er Hills at different times. When white men and on the White River, near the mouth first came, the region was in the posses- of Wounded Knee Creek, as early as sion of the Teton- Dakotas, more com- 1828. called the Sioux. the monly However, Scientific exploration of the region Sioux were immigrants, drifting west began in 1852, when Dr. John Evans from the Great Lakes country and dis- mapped the Badlands and the eastern the as late as the placing Cheyennes foothills under the auspices of the Da\'id mid-eighteenth century. Dale Owens Geological Survey. In The Black Hills region is rich in March 1853, Congress appropriated history, much of which has been re- funds for a survey of xarious proposed corded in accounts of the pioneers that routes, along \\'hich a railroad might be settled there. I have not attempted to constructed from the Mississippi River consult the vast historical literature, but to the Pacific Ocean. Survey parties were rather have restricted myself to accounts organized by the War Department, and of the various scientific and military ex- supplies for collection of natural history plorations of the study area. Results of objects were pro\'ided by the Smith- these expeditions are recorded in several sonian Institution. The first known speci- Reports of the Secretary of War in the mens of mammals from the Hills were TURNER: MAMMALS OF THE BLACK HILLS taken b\ these expeditions. Topographi- cable to the Laramie Mountains (Wade, cal and geographical knowledge of the 1947:393). country along thc> White and Bad rivers Hayden, with Captain W. F. Ray- was incremented in 1855 with the mili- nolds of the Yellowstone Expedition, Wil- tary expedition headed by General passed through the northern Black Hills Ham S. Harney, and including topograph- in 1859, reaching Devil's Tower on 20 ical Lieutenant G. K. Warren, 1868 . Both the eastern engineer July ( Raynolds, ) This and geologist Dr. F. V. Hayden. and western margins of the Hills were party passed along the southern periph- reconnoitered by elements of the Pow- ery of the Black Hills from Fort Laramie der River Expedition of 1865. In spite enroute to Fort Pierre (Warren, 1856). of the pressure to open up the Hills for accom- Two years later, Hayden again exploitation of gold, the Indian Treaty panied Lt. Warren to the region, record- of the Peace Council of Laramie, signed ing observations on 44 kinds of mam- on 29 April 1868, gave that part of the mals (Hayden, 1859). Entering from Dakota Territory lying west of the Mis- the south by way of Stockade Beaver souri River to the Sioux, temporarily clos- Creek (Weston County), they traveled ing the frontier. the western of the Hills to along edge Increasing rumors of gold in the Kara Crook Inyan Mountain, County, Black Hills and continuing troubles with Here were Wyoming (Fig. 2). they the Indians resulted in the expedition of turned back bands of by threatening Brevet Major General George A. Custer and Sioux; Hunkpapa Miniconjou they in 1874 (O'Hara, 1929; Jackson, 1966). tra\'eled southeastward to the of vicinity Members of this expedition traveled then north to and Rapid City, Sturgis about 600 miles in 60 days, and returned Bear Butte and returned to Missouri via with numerous photographs, many ob- the River 1856; 1859; Niobrara (Warren, servations on the natural history of the 1856; (1862: Hayden, 1859). Hayden region, and gold from French Creek, summarized his observations 138-151) Custer County. Dr. Othniel C. Marsh, on the natural of the Mis- history Upper the noted paleontologist, was invited to from 1854 to souri based on excursions accompany Custer, but instead sent along 1862 on 59 kinds of mam- commenting his young assistant, George Bird Grin- from the mals, including representatives nell, who took notes on 34 species of Black Hills. mammals (Grinnell, 1875:79-84). The Some reports of travel through the expedition entered the Hills from the Black Hills in the mid-nineteenth century north, traveling down Castle Creek, must be viewed with skepticism, because south to the Cheyenne River. It re- in those days the term "Black Hills" also turned northward to the area around applied to the Laramie Range along the Harvey Peak, camping at French Creek North Platte River in Wyoming. For near the present town of Custer, and example, in 1856, Lieutenant F. T. Bryan then left the Black Hills by way of Box- and naturalist W. S. Wood were part of elder Creek and Bear Butte. Ludlow that was instructed to an excellent of the an expedition ( 1875 ) gave report build a road from Fort Riley, Kansas, to reconnaisance, and included two foldout Bridger's Pass, Wyoming (Bryan, 1858). maps of the routes of Warren in 1857, Although they did not enter the Black Raynolds in 1858, and Custer in 1874. Hills, some of the specimens of mammals The Newton-Jenney U. S. Geological Sur- collected by Wood bear this locality on vey of 1875, with Colonel Richard I. the labels. Many books and articles have Dodge commanding the military escort, been published concerning the Oregon spent five months in the Hills studying Trail of Francis Parkman in 1846; the the geology and natural resources of the numerous references to the Black Hills area (Jenney, 1868; Newton and Jenney, ill C. G. New- in these publications actually are appli- 1880). Due to health, 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

of berry, the naturalist assigned to the ex- Hills from mid-May to mid-June 1910, P. G. pedition, was compelled to resign at Fort and Moulthrop and W. Phillips Laramie on the eve of departure; thus from the Cleveland Museum of Natural 15 kinds collected in the same area in the only records of mammals ( ) History observed or taken on the trip were those August 1929 (Bole, 1935; Moulthrop, recorded bv Colonel Dodge (1876: 120- 1936). Victor H. Cahalane (1948, 1951), Wildlife Division 123, 128-i;34). as Acting Chief of the As the Black Hills opened to white at Wind Cave National Park, compiled a area in settlement, biological exploration intensi- partial list of mammals of that fied. Wetmore (1968:215-216), and Pet- the course of field work from 15 August 1935 to 10 1936 file at tingill and Whitney (1965:16-17) have February (on reviewed botanical and ornithological in- Wind Cave National Park). Somewhat A. Stebler L. R. vestigations, respectively. In the sum- later, M. (1939) and left the Dice 1939 from the of mer of 1894, Walter W. Granger ( ) University paleontological expedition sponsored by Michigan Museum of Zoology, initiated the American Museum of Natural His- field investigations that resulted in pub- the tory to the Badlands of South Dakota. lished reports on mammals from Hills. A. also He journeyed west to the Black Hills Black J. King (1951, 1955), and collected 22 species of mammals, from the University of Michigan, worked mostly from the southeastern and central in the central part of the Hills and at National Park at intervals sections (J. A. Allen, 1895a), including Wind Cave C. B. several undescribed kinds (J. A. Allen, from 1945 to 1952, and Koford 1958 the Museum of Vertebrate , from 1894a, 1894b, 1895b). From 1899 to ( ) 1911, Henry Behrens made a collection Zoology, University of California, Berke- of 30 kinds of mammals on his ranch ley, carried out investigations in the along Spring Creek, and along the foot- Wyoming sector, principally in the \ icin- hills south of Rapid City. Fifty-five of ity of Devils Tower. these specimens now are housed in the In 1947, 1951, 1961, 1965, and again in Pioneer Room of the Minnilusa Histori- 1967, field parties from the Museum of cal Museum in Rapid City, South Da- Natural History of The University of kota. Kansas collected mammals in western The U. S. Biological Survey became South Dakota and northeastern Wyo- active in the region at the turn of the ming (Jones and Packard, 1958; Long, century (Merriam, 1888, 1889, 1891). 1965; Jones and Genoways, 1967a, 1967b; Vernon Bailey and Merritt Gary were re- Turner and Jones, 1968; Turner and sponsible for obtaining mammals and Davis, 1970; White, 1952, 1953a, 1953b). natural historv data from the region. Be- My own work in the Black Hills began tween 1899 and 1912, Cary (1917) peri- in the summer of 1965; I returned to the odically investigated the area along the study area throughout the summers of South Dakota-Wyoming border in the 1967 and 1968. A week was spent in vicinity of Elk Mountain. Bailey (1914) quest of hibernating bats in late Novem- intermittently collected in the Black Hills ber of 1967, and a week each in obtain- from 1887 until 1913; observations of 25 ing photographs and samples of soil in species of mammals were made on his August of 1969, and March of 1970. In initial excursion near Rapid City and all, I was in the field for 31 weeks from Deadwood late in 1887 (Bailey, 1888). mid-June 1965 to March 1970, including The materials, gathered by Granger, two months as a Ranger-Naturalist at Bailey and Gary contributed significantly Wind Gave National Park in 1968. Ad- to the present study. ditionally, tvvo weeks were required in to from the A. H. Howell, N. Dearborn, and a 1968 examine specimens other field party from the U. S. National Mu- Black Hills housed in museums seum worked in the central part of the than at Kansas. TURNER: xMAMMALS OF THE BLACK HILLS 11

METHODS AND MATERIALS for taxonomic clarification or when con- variation is evident. Each of the six orders of Black Hills spicuous geographic 6. All measurements, external and mammals and the 19 families ( discussed are in millimeters. Stand- briefly), including 44 genera and 62 cranial, given ard external measurements were read species, are arranged in text following from labels attached to the arrangement of Hall and Kelson the specimens, of the (1959). The species of each genus and, excepting length forearm; latter is to bats and was meas- where appropriate, the subspecies of applicable only ured from museum each species, are in alphabetic order. prepared specimens. of adult males and Keys to species, disti-ibution maps, and Weights non-pregnant adult females are in Stated enumeration of characters applicable to given grams. measurements are the arithmetic the \arious ta.xonomic categories, are not mean, followed the standard deviation. Un- pro\ ided herein as these were deemed by less otherwise measurements are to be readily accessible in other publi- stated, cations. of adults only. Cranial measurements were taken to the nearest tenth of a The account of each monotypic spe- millimeter with dial in the man- cies or subspecies native to the Black calipers ner described Hall Hills incorporates the following: by (1946:672-685), and Packard identical 1. The scientific name, employed in (1960:584-585); agreement with the International Rules measurements were not taken for all of Zoological Nomenclature, and fol- species. lowed on the same line by the name of Variation in color was assessed by the author. direct comparison of specimens and also 2. The vernacular name, in general by use of a Photo\'olt Photoelectric Re- accord with Hall 1965 that flection Model ( ) , and which Meter, 610, utilizing red, is considered appropriate for all sub- green, and blue filters. Readings were species of the given species. taken from the middorsal region of mu- 3. The synonymy, in which the first seum study skins and could be repeated citation is to the original description and on the same individual with minimal is followed by designation of the type variation. Samples of soil from various locality. The second citation is to the collecting sites were first dried for sev- first usage of the currently accepted eral hours in an oven, then subjected to name-combination employed herein, un- color analysis with the reflection meter. less that combination is identical to the Resultant measurements of reflected name as originally proposed. In a few light were recorded for each skin and cases, a third citation is to a taxon de- soil sample as percentage values of pure scribed from the Black Hills, but now white calibrated against a standardized placed in synonymy. block of magnesium carbonate. These 4. The disirihuiion, which concerns recorded measurements actually repre- the regional distribution of each species sent a composite value of both intensity and comments pertaining to the abun- of hue (that is, more red or less red in dance and apportionment of each color) and tone (paler or darker in throughout the Black Hills. Included color ) . For example, pale brownish soils here are notations on ecological, altitudi- yielded higher reflectance readings when nal, and seasonal distributions; the latter using a red filter than did dark reddish is especially important for those mam- soils. Reflectance readings obtained from mals that hibernate or migrate. all three filters were summed in order to 5. Comments on systematics are in- achieve an approximate index of tone. cluded in most accounts. Comparisons If, for instance, the specimens or soil of characters are made between the samples were arranged in order of de- Black Hills populations and those of sur- creasing paleness of overall coloration, rounding areas when these are warranted the resultant sequence derived by direct 12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY comparison would be in agreement with 8. The records of occurrence, include the summed total (trichromatic) reflect- both specimens examined (based on 4727 additional records. ance, or tone (Figs. 13 and 14). Inten- specimens) and sity of each hue was derived by dividing Under specimens exatnined, the first the respective initial reflectance reading notation is the total number of speci- from each filter by the total reflectance mens examined by me, followed by the readings from all three filters. Extremely exact locality of capture, the number of skins were not in- greasy or damaged specimens from each locality, and desig- cluded in the color analysis. nation of site of specimen deposition. In analysis of geographic variation, Localities are allocated to their respec- specimens were segregated by age, sex, tive counties, which are grouped under season, and geogi'aphic origin. The either South Dakota or Wyoming. measurements obtained under these County names and the localities within groupings were then subjected to stand- each county are arranged from north to ard univariate statistical analysis and to south (and \\est to east if more than Variance. If the an overall Analysis of one locality occurs in the same latitude Analysis of Variance indicated that sig- within a county). The many specimens nificant differences existed among examined for comparisons from areas groups, then the Sums of Squares Simul- surrounding the Black Hills are not enu- taneous Test Procedure of Gabriel merated. Abbreviations designating (1964) was applied in order to locate specimens examined in collections other these differences. than the Museum of Natural History of 7. Remarks on autecoIog,i/ ordinarily The University of Kansas are as follows: include information pertinent to the nat- AMNH—American Museum of Natural ural history of each species. Methods of History, New York; FMNH—Field Mu- capture, time of activity (i.e.. Mountain seum of Natural History, Chicago; MHM Daylight Time, MDT, or Mountain —The Pioneer Room, Minnilusa Histori- Standard Time, MST), description of cal Museum (collection of Henry Beh- habitat, enumeration of associates, and rens Rapid City; MSB—Museum of additional noteworthy observations are Southwestern Biology, University of recorded. A brief chronicle is given for New Mexico, Albuquerque; NRW—Col- those species that were common in his- lection of N. R. Whitney, Rapid City; toric times, but now have been reduced SDMT—Department of Biology, South in numbers or extirpated. Reproductive Dakota School of Mines and Technology, data are summarized for each species for Rapid City; SDSU—Department of En- which information (usually taken from tomology and Zoology, South Dakota specimen labels or field notes of the State University, Brookings; UK—De- collector) is available. Molt data were partment of Zoology, University of Ken- obtained from examination of museum tucky, Lexington; UMMZ—Museum of specimens, both diy and in alcohol, by Zoology, University of Michigan (in- directing a stream of air through the cluding specimens formerly housed in pelage and noting the presence or ab- the Cleveland Museum of Natural His- hairs. In recent sence of underlying new tory), Ann Arbor; USD—Department of year, specimens were examined in the Biology, University of South Dakota, field immediately after capture for ecto- Vermillion; USNM—United States Na- Parasites collected in this parasites. tional Museum (including collections of manner were preserved in 70 percent the U. S. Biological Surveys), Washing- alcohol and referred to various special- ton, D. C; UW—Department of Zoology, ists for identification. In compiling the Uni\'ersity of Wyoming, Laramie; above-mentioned data and observations, Collection of Wind Cave Na- field notes of 46 individuals were con- WCNP— tional Hot sulted, in addition to my own firsthand Park, Springs. observations. Additional records consist of reports TURNER: MAMMALS OF THE BLACK HILLS 13 from the literature, specimens in collec- of B. Bailey that were identified by per- tions that I have not examined, or identi- sonnel of the Survey. Localities from fications of presumed accuracy as re- which specimens were examined are not corded in card files of the U. S. Biologi- duplicated in the additional records. cal Sur\'cy (USES); the abbre\'iation Most sites from which mammals were BB designates specimens in the collection obtained are shown in figure 2. ENVIRONMENT

GEOGRAPHY ogy of the Black Hills. The reader is directed to Darton (1909), Darton and The Black Hills constitute a maturely Paige (1925), Mcintosh (1931), Tullis dissected, isolated, mountainous region (1951), and Cries and TulHs for of approximateh' 4000 square miles that (1955) detailed geological accounts of the re- resulted from intermittent domal uplifts in Cretaceous, Miocene, and Pleistocene gion. times. The area extends 120 miles in a The Black Hills uplift is a crescent- northwest-southeast direction, and is 50 shaped, anticlinal dome that rises sev- eral thousand feet above the miles wide at the widest point. The Hills surrounding Northern Great Plains 1 and are entirely surrounded by the non-glaci- (Figs. 3). ated Missouri Plateau section of the The Central Basin is located slightly east of the main axis of the dome. It is Northern Great Plains physiographic a mountainous core of province (Fenneman, 1931), and rise rugged Pre-Cam- brian rock and abo\e the plains to an elevation of about igneous Pre-Cambrian, 4000 feet on the east and 3000 feet on Paleozoic and Mesozoic sediments. The Basin is with the west. The highest point (Harney interspersed park-like val- and Peak) lies 7242 feet above sea level. leys steep, narrow canyons that in- cline to the and east. In the Most of the region lies within the drain- north, south, Peak-Needles-Mount age of the Cheyenne River, which circles Harney Rushmore area 4 there are bare the south end of the Black Hills, and ( Fig. ) granitic and the the Belle Fourche Ri\'er, which skirts mctamorphic ridges. Encircling Cen- tral Basin is the Limestone Plateau of the north edge of the area (Figs. 1 and 5 and 2). rolling highlands (Figs. 10). The is about two miles in width to As defined herein, the Black Hills are plateau the east, and reaches a maximum of 15 delimited by the distribution of Jurassic shale and sandstone of the Sundance miles in width in the northwestern por- tion of the Black Hills. It is Formation 3 . the comprised ( Fig. ) Thus, region of Paleozoic sediments shale includes the Black Hills proper, as well (sandstone, and limestone of the Deadwood, White- as the Bear Lodge Mountains and Devils ) wood, and Minne- Tower of Wyoming; the latter two areas Pahasapa-Englcwood lusa formations. The wall-like are closely allied with the Hills. Ranges escarp- ment of the reaches elevations of mountains nearest the Black Hills are plateau of 7100 feet; it faces inward toward the the Laramie Mountains (elevations to Central Basin and is interrupted such 10,272 ft) and Big Horn Mountains (ele- by stream valleys as Sand, Cold Spring, and vations to 13,165 ft); these lie approxi- creeks. In some areas mately 150 miles to the southwest and Spearfish (e.g., and Little west, respectively, in Wyoming. Big Spearfish Canyons; Fig. 6) of the Limestone Plateau, high cliffs pro- GEOLOGY AND PHYSIOGRAPHY ject above the valley floors. The rugged terrain in the northern Central Basin and As a result of the presence of a di- northwestern Limestone versity of geological structures and pro- Plateau was ductive mineral deposits, there are nu- formed by solidification of igneous in- merous publications concerning the geol- trusions during Tertiary time (Fig. 3). 14 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

I0 4°

44< 44<

I04°

Fig. 1. General physiography of the Black Hills of South Dakota and Wyoming (modified after A. N. Strahler, Physical Geography, John Wiley and Sons). Encircled numbers indicate sites at which photographs of the corresponding figures were obtained.

Sloping outward from the plateau below). The foothills are less extensive a are broadly rolling tablelands that com- along the eastern slope due to steeper prise the foothills and the Foothill Tran- gradient, and are composed of Paleozoic limestone and sition Soil Association ( see SOILS section (Minnekahta Formation) TURNER: MAMMALS OF THE BLACK HILLS 15

104'

_ PIACERVIILE PACIOLA '/ \_^ C_N_^_ ""

f

NIOBRARA CO.

ANGOSTURA RESERVOIR 20 EDGEMONT Hl MILES

104°

Fig. 2. General geography of the Black Hills, showing major collecting sites mentioned in te.\t (modified after Pettingill and Whitney, 1965:frontis). 16 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

104°

104"

15J Tertiary igneous intrusions Paleozoic sediments

I I Tertiary sediments 'w'.l Precambrlan granitic rocks

. Cretaceous sediments Precambrlan metamorphic rocks

Triassic and Jurassic sediments

Fig. 3. Generalized geologic map of the Black Hills area (modified after Black Hills Area Resource Study, 1967:18). TURNER: MAMMALS OF THE BLACK HILLS 17 shale (Opcche Formation). Along the abundant on the Limestone Plateau than base of the foothills lies a conspicuous, elsewhere in the Hills. Because of the and remarkabK' continuous valley in the plateau's broad valley meadows and form of a "racetrack," \\'hich encircles the proximity to the foothills, it also provides Hills (Fig. 7). This narrow trough rep- optimal habitat for both the mule deer resents the lowest elevation in the Hills (Odocoileus hemionus) and white-tailed

(3200 to 3500 feet above sea level). It deer (O. virginianus) . Streams transect cuts into soft red shales and sandstone of all of these areas to produce the Un- the Spearfish Formation, which was differentiated Alluvial Soil Association formed as a deposition of sediments that is inhabited by various riparian spe- early in the Triassic. The Red Valley, so cies such as the masked shrew (Sore.t named because of reddish soils of the cinereus) and meadow jumping mouse

Spearfish-Ne\'ille Soils Subassociation, {Zapus liudsonius) . varies in width from a of a mile quarter Faunas of two isolated Tertiary ig- the to more than six miles (in east) (in neous protrusions are noteworthy. Inyan the is dis- the northwest); xalley floor Kara Mountain, surrounded by foothills sected divides that are by many drainage and upland prairie in southeastern Crook broad alluvial flats. separated by County, Wyoming, is the primary range The Limestone Plateau, Central Ba- for a herd of recently introduced moun- sin, and peaks of the Bear Lodge Moun- tain sheep (OvLs canadensis) a species tains form the so-called "boreal-cap" of formerly native to the Hills (Baird, 1858: the Black Hills; the latter two areas 678; Cowan, 1940:543). Devils Tower is comprise the Mountainland Soil Asso- an igneous monolith that proti-udes 865 ciation, whereas the former consists of feet above the surrounding prairie and the Limestone Plateau Soil Association. open pine woodland. The fissures and The boreal-cap is characterized by a pre- surface atop the Tower are well carpeted dominantly coniferous forest fauna of with lichens, grass, sagebrush, and prick- boreomontane and cordilleran origin. lypear cactus, and support a meager Mammals such as Nuttall's cottontail fauna consisting of least chipmunks, deer (Sylvilagus nuttallii), the least chipmunk mice, and bushy-tailed woodrats {Neo- {Eutamias minimus), yellow-bellied toma cinerea) (exhibit. Visitor's Center, marmot (Marmota flaviventris), red Devils Tower National Monument). squirrel (Taniiasciunis hudsonicus), Higher areas in the foothills support sa- northern flying squirrel (Glaticomys a fauna that is similar to that of the northern ])rinus), pocket gopher {Tho- boreal-cap, whereas the fauna of the red-backed vole momys talpoides), lower regions more closely approaches and (CletJirionomys gapperi), long- that of the surrounding prairies. For tailed \'ole Microtus reach ( Jongicaiidtis), example, along the foothill-Limestone their highest population densities in these Plateau border, the meadow vole often areas. In the Central isolated Basin, is captured in the same trapline with the of the Slate Prairie Soil Asso- grasslands long-tailed vole. Along the foothill-Red ciation 11 are inhabited ( Fig. ) mainly by Vallev transition, the meadow vole is the white-tailed toicn- jackrabbit (Lepus associated with the prairie vole {Micro- semlii), thirteen-lined squirrel tus and the ground ochrogaster) , long-tailed deer {Spennophihis tridecemlineatus) , vole is not present. Typical mammals of mouse and {Peromysctis maniculatus) , the Red Valley grasslands and lower meadow vole {Microtus pennsylvani- foothills are the desert cottontail {Syl- The core near cus). crystalline Harney vilagus audubonii), white-tailed jack- Peak the restricted of the provides range rabbit, thirteen-lined ground squiiTcl, introduced mountain goat (Oreamnos black-tailed prairie dog {Cynomys liido- americanus). Both Nuttall's cottontail vicianus), deer mouse, western harvest and the white-tailed jackrabbit are more mouse {Reithrodontomys megalotis). 18 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Fig. 4. The Needles, exposed crystalline rock in the Mount Rushmore-Hamey Peak area of the Central Basin. These moss- and lichen-covered granitic ridges thrust upward nearly perpen- dicular to the surrounding forest, and comprise the restricted range of the introduced mountain goat.

Fig. 5. A broad upland valley on the rolling slopes of the Limestone Plateau, south of Moon. Such areas generally are farmed in oats, legumes and tame grasses, or are grazed, but also support large herds of deer. TURNER: MAMMALS OF THE BLACK HILLS 19

t "^F? ml/'i»»i

Fig. 6. Due to resistence to erosion, high cliffs project as steep walls above canyon floors in more rugged areas of the Limestone Plateau, such as Little Spearfsh Canyon. The above photograph was taken from the rear entrance of a cave, in a cliff above the Timon Camp- ground, that served as a well-used night roost for several kinds of bats (see account of small-

footed myotis ) .

Fig. 7. The Red \ alley, locally termed the "racetrack," is a remarkably continuous trough that encircles the Black Hills, and occupies a valley-like position between the footliills and the Dakota Hogback. Large herds of bison, elk, and pronghom, and colonies of prairie dogs inhabit the extensive grasslands comprising the Red Valley portion of Wind Cave National Park. 20 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

^'^:-j^£n\'

Fig. 8. The outer border of the Red Valley, and of tlie Black Hills, is formed by a steep- sided, 600-foot sandstone ridge, termed the Dakota Hogback, that slopes gradually outward onto the surrounding semiarid mixed grass prairie. The above break in the ridge (Buffalo Gap) has been eroded away l)y Beaver Creek as it flows out of the Black Hills. Large herds of bison and pronghorn previously migrated between the grasslands of the Red Valley and those of the Northern Great Plains through this stream gap, which also pro\ided access for Jedediah Smith in 1823 and later explorers and adventurers.

Fig. 9. The predominant vegetational component of the Black Hills, especially in the Central Basin, is a montane belt of ponderosa pine. This conifer seems tolerant of xerophytic condi- tions and grows even on exposed rocky ridges. The dark appearance of the foliage of the ponderosa pine, when viewed from a distance, accounts for the Sioux Indian name Paha Sapa, or "Black Hills." TURNER: MAMMALS OF THE BLACK HILLS 21

Fig. 10. Moist, narrow valleys in the Limestone Plateau support a dense subalpine belt of white spruce with an associated understory of northern affinities. Luxuriant riparian habitats such as that bordering the Beaver Creek Campground are occupied by masked shrews, meadow jumping mice, and several kinds of voles.

:^ -*

9^- ^ •% .

Fig. 11. The Bald Hills is one of three isolated grasslands in the Central Basin situated on ^\ell-rounded slopes underlain by slates. There is no evidence tliat these grasslands have ever been forested (Photograph by Barry L. Siler). 22 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Taxidea taxiis and It was this stream that prairie vole, badger ( ) , through gap large coyote (Canis latrans). In addition to herds of bison (Bison bison) previously those species just mentioned, the Sandhill migrated into the grasslands of the Red Regosol Soil Subassociation along the Valley. Custer County, South Dakota-Weston CLIMATE County, Wyoming border is inhabited by se\'eral heteromyids. Included among The climate of the Black Hills is dis- are the the latter hispid pocket mouse tinct from that of the surrounding, semi- (Perognathus Impidiis), oHve-backed arid Great Plains in being more moist and Ord's pocket mouse (P. fasciatus), and less subject to extremes of tempera- kangaroo rat {Dipodomys ordii); all are ture. Because of warm "chinook" winds characteristic of rocky or sandy soils. and frequent sunny skies, the Black Hills The outer border of the Red Valley are the warmest part of South Dakota in is formed by a 600 foot sandstone rim winter. An additional moderating influ- that is steep-faced within, but slopes ence is the tendency for hea\y colder air gradually outward onto the surrounding to seek low elevations; thus, Arctic air Great Plains (Fig. 8). This hogback masses that blanket the plains in colder ranges in e]e\'ation from 3800 to 4900 winter months bypass the higher eleva- feet (near Elk Mountain). The outward- tions of the Hills. Conversely, summer sloping cuestas frequently are broken by temperatures at higher elevations in the streams. The inner face of the hogback Hills are cooler than are those of the is of Jurassic age and is composed of surrounding prairies. green shales and red sandstones of the Contrasts in climate between Lead, Sundance Formation and to a lesser ex- in the north-central part of the Black tent of the shales, sandstones, and lime- Hills, and Hot Springs, at the southern stones of the Morrison Formation. The periphery, are shown in table 1 and fig- outer slope is judged as Cretaceous in ure 12. Average maximum temperatures age and formed of the Skull Creek and for July range from 81-83' F in the Black Inyan Kara formations, both of which are Hills and 88-92 F on the adjacent plains; composites of Dakota and Lakota sand- axerage minimum temperatures for Janu- stones, Fuson shale, and Minnewaste ary range from 9-14^ F in the Hills and limestone formations. The conspicuous 4-9° F on the surrounding grasslands. hogback, marking the boundary between Warm days and cool nights are charac- the Black Hills and the Northern Great teristic of the Hills. Air temperatures Plains, forms a series of ridges that were usually have a daily range from 50-60° F an obstacle to early travelers. Beaver in the shade in summer, but the range Creek has eroded away a break in the may reach 80°, or more, in direct sun- hogback, just northwest of Buffalo Gap. light (Wetmore, 1968:217).

Table 1.—Temperatiue and precipitation data for Hot Springs (southeastern Black Hills) and Lead (north-central Black Hills), 1931-1935 (Hodge, 1960:8). TURNER: MAMMALS OF THE BLACK HILLS 23

80

'j^

60-

40- zH- o z < LU ^ 20

^ -f 2 4 MEAN MONTHLY PRECIPITATION (INCHES)

Fig. 12. Hydrothermograph for Lead (solid line) in the north-central Black Hills, and Hot Springs

( broken line) in the southeastern Black Hills, 1931-1935 (Hodge, 1960:8). Labeled circles ( X ) indicate annual means of both parameters for each climatological station.

The frost-free season is shortest at tion. Hailstorms generally occur in mid- higher elevations, where brief freezing summer, and lightning usually in late has been known to occur at any time of summer. summer; however, there are usually only 110 frost-free days in the Black Hills as SOILS compared to 130 on the pheripheral Materials from which soils have de- prairies. veloped in the Black Hills include an- Average annual precipitation in the cient crystalline rock in the central part Black Hills ranges from 16 inches, in the of the region and rocks southern portion, to 28 inches in the sedimentary ( shale, sandstone, and limestone in out- northern section, where snow and rain ) lying areas (Figs. 1 and 3). Local topog- often are formed when prevailing winds raphy contributes to soil formation by are forced abruptly up steep sides of determining drainage; thus, steep slopes mountains. Mean annual precipitation have well-drained, thin soils, whereas on the adjacent plains is 14-16 inches. more level areas have more poorly Much of the total precipitation occurs drained, soils. as rain during summer in the drier south- deep The soils of the Black Hills are ern portion of the Hills, whereas it is major much more c\'enly distributed through- the Gray Wooded Soils; they are unique cut the year in the moister northern sec- to the general region because they de- 24 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY veloped under forest in a relatively hn- horizon is organic and about two to four mid climate. Soils of the surrounding inches in depth. The B- horizon is be- area developed under grasslands in a tween 10 and 15 inches in depth and of climate ranging from moist subhumid to prismatic structure with clayfilms on semiarid. The horizons of Gray Wooded most surfaces. The B.-; horizon is at least Soils of the Black Hills are subdivided six inches in depth and structured of as follows: 1) Aon (forest litter). Am prisms. Both the B.. and C horizons (partly decayed litter), and A02 (well contain free carbonates. decayed litter); these are horizons which Azonal soil groups in the Black Hills are organic and thin (i.e., one-half to two include Regosols, Lithosols, and Allu- inches in depth). 2) Ai —a mineral hori- vium. Regosols are thin soils developed zon that is thin or absent. 3) A:>—a hori- in unconsolidated material, whereas zon that is weakly platy in structure and Lithosols are thin soils over solid rock from four to 12 inches in depth. 4) Bi —a (within 1(S inches of the surface). These four-inch-thick transitional layer between immature soils occupy steep slopes where the Ao and B2 horizons; Bi is absent in runoff is so excessive that little leaching the Hills. 5) B-—a horizon which is or development of humus occurs. No brown and blocky in structure and pos- true subsoil is present. Alknium occu- sesses clayfilms on all surfaces; this layer pies flood plains of various water courses is between 10 and 30 inches in depth. in the Hills. Alluvium tends to be darker 6) C—a horizon which varies in compo- and more humic than soil of the adjacent sition in different areas. uplands and frequently supports a not- Other zonal soils that occur on lower ably different riparian plant community. sloDes are the Chernozems and Chestnut A great need exists for detailed infor- soils. Color of the surface layer of the mation on soils in the Black Hills. Most Chernozems is black to dark, grayish of the soils are unclassified and are stony brown. Soils of this type occupy open or rocky, reflecting the local geology and woodlands and are paler in color and physiography. For detailed information browner in the southern part of the Hills concerning land use of the soil associa- than in the northern sections. Ai hori- tions, the reader is directed to Austin zons usually are five to eight inches in (1965:26-28), Westin ef al. (1967:1-32) depth and granular in structure. Bo and to miscellaneous publications issued horizons arc about eight to 10 inches in by the Soil and Water Conservation Dis- depth and structured of prisms that ordi- tricts of the respective counties. Some narily are coated with clayfilms on all relationships of soils to the distribution surfaces. B.^ horizons have a weak, pris- and speciation of mammals in the Black matic structure; they are usually more Hills are discussed bevond. than six inches in depth and more clayey The following description of Soil As- in composition. Both the B... and C hori- sociations in the Black Hills (referred to zons are enriched with lime leached from in text) has been synthesized from frag- the upper layers. The surface layer of mentary literature and in consultation Chestnut soil is brown or reddish in color with the South Dakota Soils and Water and developed under grasslands. The A, Conservation Service.

Mountainland Association.—occupies the Central Basin mountainous core (1200 square miles) of igneous and sedimentary rock; excessively drained; topography: high rocky ridges, narrow loll- ing plateaus, deeply entrenched canyons, and park-like mountain valleys; exposed slopes: bare rock; steep slopes: Spearfish and Laporte Lithosols; upper forested slopes: Edloe Gray Wooded Soils; intermediate woodland slopes: Chernozem Soils; lower grassland slopes: Chest- nut Soils; along valley and canyon drainages: -Table Mountain Soils (alhnial loams, inter- mixed with loose stones); land use: recreation, major timber production (ponderosa pine), and grazing. Limestone Plateau Association. —occupies a high plateau of sedimentary rock (sandstone, lime- stone and shale) that encircles the Central Basin; well-drained; topography: rolling slopes, TURNER: MAMMALS OF THE BLACK HILLS 25

broad upland valleys, few rocky ridges, buttes and steep canyons; distribution of major soil types as in the Mountainland Association, differing only in percent coverage due to contrasts in topograph)'; rocky silt loams that occup\' ridges may be absent on south-facing slopes where stands of pine are thin or absent; grassland soils of some upland \'alleys have a water table; land use: recreation, some timber production (ponderosa pine and white spruce), grazing, and farming (oats, legumes and tame grasses).

Slate Prairie Associatiou.—occupies three isolated prairies (Reynolds and Gillette Prairies, and the Bald Hills) comprising about 9000 acres in the Central Basin; well-drained; topography: rolling to steepK' rounded slopes; shallow rocky loams underlain by slates, some Table Moim- tain Soils along upland drainages; no evidence of prior forestation; land use: grazing and

farming (oats and Aegetables ) . Undifferentiated Alhivial Association. —occius along stream channels of all major valley systems in the Black Hills; well-drained, except for some areas of seepage; topography: fairly level to genth' undu'ating; deep silt loams, with weak subsoil development; land use (highly pro-

ductive, but limited by cool temperatures and short growing season ) : grazing and fanning (oats, alfalfa and hay). Foothill Transition Association.—inter\'enes between the Limestone Plateau and Dakota Hogback; well-drained; topography: complexly dissected foothills with gently rounded to steep slopes, and narrow ridges and \alleys; land use: recreation, minor timber production (ponderosa pine), grazing, and dryland and irrigational farming. Due to topographical di\'ersity, this region is di\ided into several subassociations: Laporte-Sandoz-Berthoiid Suhassociation.—calcareous soils with limestone outcrops often exposed on the surface; Laporte Lithosols: occur on ridge tops and abrupt slopes; Sandoz Chernozems: occupy intermediate woodland slopes; Berthoud Chestnut Soils: occupy alluvial swales and drainageways on lower grassland slopes. Spearfish-Neville Sidmssociation.—reddish-colored Chestnut Soils imder grasslands that occupy a \alley-like ("race-track") position between the steeper foothills and Dakota Hogback and overlay a g>'psum-like shale substratinu; Spearfish Soils: occur on gently rolling slopes; Neville Soils: occupy longer and smoother colluvial and residual slopes. Sand Hill Re^.osol Suhassociation.—sands and fine sandy loams (presumably of the Valen-

tine series ) extend northw estward from the Sand Hills of Nebraska, along the Custer County, South Dakota-Weston County, Wyoming border; occur on gently rolling ter- rain; exposed rock prominent in some areas. Dakota Hogback Association.—occupies a steep sandstone ridge that encircles the outer limits of the Black Hills; well-drained; topography: abrupt to sloping, with exposed sandstone out- crops; shallow calcareous Travessila Lithosols; land use: grazing.

The Pierre Sliale Plains Association occurs just exterior to the Black Hills, consisting of grassland Chestnut Soils with moderately deep firm cUiys overlaying shales, and is penetrated by the Vale- Bcaverton Alhivial Association that occupies low terraces and benches adjacent to larger streams that flow out from the Hills. Soils of Southwestern South Dakota, excluding the Black Hills, have the palest tone, are browner in hue, and ha\ e less organic matter and total nitrogen than soils of other parts of the state.

HYDROGRAPHY River, with a surface drainage of 2000 square miles, flows the southern Description, classification, and utiliza- along these rivers have tion of lakes and streams in the Black periphery. Together, an annual flow of Hills have been discussed in the Black 522,000 acre-feet; they on the about 60 miles east Hills Area Resource Study of 1967, and join prairie of the Black Hills and flow eastward by Stewart and Thilenius (1964). All (as the into the Missouri River. water courses that arise in the Black Cheyenne) The northwest-southeast of Hills comprise 1302 miles of sh-eam alignment their major tributaries, such as Boxelder, drainage, and empty either into the Belle Rapid, Spring, Battle, French, and Fourche or Cheyenne rivers; both rivers Beaver creeks and of other streams in originate on the gently rolling plains to western with the west in Wyoming. The Belle Fourche South Dakota, coincides the wind and River, with a surface drainage of 7210 prevailing direction, ap- square miles, borders the northern mar- pears to be the result of periodic accumu- gin of the Hills; whereas, the Cheyenne lation of locally derived eolian sediments 26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

(White, 1961:207). Although June is cally reduces productivity of the lake the time of maximum precipitation, max- (Stewart and Thilenius, 1964:46). imum surface runoff is in May owing to Dodge (1876) recorded the hydrog- the greater evapotranspiration rates en- raphy of the Black Hills when he ac- countered later in the growing season. companied the Newton-Janney U. S. Floods are common in both months. Geological Survey of 1875. Stream flow was as substantial Due to abrupt topographic relief, reported and the wa- most streams in the Black Hills are ters as being "cool, clear, and pure." Al- radial-dendritic systems that flow swiftly; though riparian vegetation was suffi- only in a few montane meadows and ciently dense to impede travel along the stream broad valleys does a more gradual gra- bottoms, forest cover evidently dient allow some meandering and for- was less extensive then than now, in that mation of small, semimarsh areas ad- large older trees were "scarcely to be jacent to stream channels. In addition found." Adequate fire protection and to aquatic mammals such as the beaver management practices such as "thinning" (Castor canadensis) and muskrat (On- have allowed a mature forest with a closed datra zihethicus), riparian species such canopy to develop over much of the Hills. as the mink (Mustela vison), masked Thickening of the forest, with resultant shrew, jumping mouse, long-tailed vole, interception of precipitation and meadow vole inhabit these moist and increased evapotranspiration, is the environs. greatest single factor in reduction of total surface and in decrease When European man first entered discharge of moisture available for the Black Hills, beaver ponds were the recharging water stores and Thil- only sources of standing water. Now the ground (Stewart enius, of foothills are interrupted by thousands of 1964). Trampling riparian cattle around small reservoirs that store water for live- vegetation by springs and creeks, construction of roads stock and for fish and wildlife; major along stream channels, of wa- impoundments such as Angostura, Deer- tapping ground ter with wells, and utili- field, Pactola, Keyhole, and Belle supplies storage zation of water for Fourche reservoirs also have been con- agriculture and indus- and of structed. These artificial bodies of try, deposition tailing from mines, tannic acid from saw mills, and wastes standing water (usually located over from continue to shist or shale) are used for recreation, as municipalities degrade habitats in the Black Hills. well as for watershed development and aquatic irrigation. VEGETATION Except for the headwaters, all streams in the Black Hills are affected to some The most conspicuous component of the degree by pollution. Whitewood Creek Black Hills vegetation is the conif- erous is is the most severely polluted stream in forest, which dominated by a the Hills; it carries an extremely high montane belt of ponderosa pine (Pintis In content of rock-flour and chemical pollu- ponderosa) (Fig. 9). more mesic sites on northern tants from mining operations and a heavy exposures in the Central Ba- sin and on the load of municipal wastes from Lead and Limestone Plateau, there is a belt of white Ficea Deadwood. The North Fork of Rapid subalpine spruce ( iijauca) with an associated understory of Creek also is polluted with a high con- plants with northern affinities (Fig. iO). tent of bog-iron wastes. Stockade Lake Kentucky bluegrass (Poa prafensis), in- is of poor equality due to sewage effluents troduced from Europe in the 17th Cen- from Custer. A dense bloom of a toxic tury, is the predominant graminoid plant blue-green algae (Aphanizomenon flos- of the open meadows, parklike areas, and results from aqiiae) the supplemental grasslands of the foothills and Red Val- and nitrogen phosphorous, and drasti- ley. The flora is a mixture of boreal. TURNER: MAMMALS OF THE BLACK HILLS 27

cordillrran, eastern deciduous forest, and cedar (Jiiniperus virginiana) intrudes Great Plains species (Rydberg, 1896; among the pine. Along the cooler canyon Webb, 1965). Biogeographic affinities of floors and northern slopes of the central the Black Hills biota ha\e been treated and moist northern sections of the Black hv \arious authors. Birds are discussed Hills, the coniferous forest includes white bv Miller (1941), Pettingill and Whit- spruce, paper birch (Betida papyifera), ney (1965), and Mengel (1970). Byers and quaking aspen (Populus tremu-

(1961) and Ross (1965; 1970) dealt with Joides) . The accompanying understory 1957 the nti- insects, and Smith ( ) discussed consists of wild sarsaparilla (Aralia smooth green snake. Willo\\'s and lichens (Ucoidis), twin-flower (Linnaea hore- were treated Froiland 1962 and currant by ( ) cdis), swamp (Rd)es Jactistre), \\'etmore Forest red (1968), respectively. bunchberry (Cornus canadetisis) , and other are osier Venus' vegetational components (C. stolonifera) , slipper discussed Buttrick 1914 by ( ) , Halliday (Calypso hidhosa), squashberry (Vi- and Brown (1943), Havward (1928), hurnum edide) buffaloberry (Shep- Mcintosh (1931), Dillon' (1956), Potter Jierdia argentea), and huckleberry (Vac- and Green (1964), and Watts and cinium memhranaceum) in close asso- Wright (1966). ciation. White spruce may occasionally The most exposed sections in the dominate the coniferous forest of lower in these but it oc- Black Hills, such as the crown of Harney slopes areas, usually Peak and the Needles section, either are curs in subalpine belts that are in close to montane belts of de\'oid of ^'egetation or support bryoids proximity ponderosa Predominant associates (mosses) and thallophytes (lichens). A pine (Fig. 10). of the are various climatic timberline is not present; in- pine bunch grasses, stead, the absence of trees along the bearberry [ArctostaphyJos uva-ursi), high summits is due to lack of soil and wild rose (Rosa acictdaris), creeping to extreme exposure. In general, three juniper (Juniperus horizontalis) , ground vegetational types occur in the Black juniper (/. communis), Oregon grape Hills—coniferous forest, deciduous (Berlyeris repens), redroot (Ceanothus tea woodlands, and prairie grasslands. Dry velutintis) , New Jersey (C. ovatus),

and Rhus radicans . slopes are dominated by ponderosa pine, poison ivy ( ) the dark appearance of which accounts Deciduous Woodland Association.— for the Sioux name PaJia Sapa or "Black Bur oak (Que reus macrocarpa) com- Hills." Usually broadleafed trees are de- monl\' is intermixed with the pines along veloped mainly along drainage channels, drainages near the periphery of the Hills; or are present as groves on old, burned it occurs in drier valleys, on drier slopes, areas. Grasslands are found mostly on and on sandy to gravelly soils. Quaking slopes of the foothills, but also in isolated aspen, paper birch, green ash (Fraxinus areas even at elevations. prairie higher pennsylvanica) , and American elm (Ul- Coniferous Forest Association.—This nius americana) are also present. In the association forms a forest that is domi- northern section, and especially in the Bear is the nated by ponderosa pine, which appears Lodge Mountains, bur oak to be extremely tolerant of xerophytic dominant tree growth on many slopes stream channels. The conditions (Fig. 9). The species ranges leading away from between elevations of approximately oak has an undergrowth of hawthorn

3500 and 7000 feet, and forms parklike (Crataegus chrysocarpa) , chokecherry forests at higher elevations that give way (Pyrus virginiana), hop hornbeam (Os- and to open woodlands at lower elevations. trya virginiana) , poison ivy. On the pine-clad uplands, the soil is Riparian Association.—Streamside coarse, well drained, and quite warm in habitats in the Black Hills are character- the summer. In the western section, and ized by fluviatile soils that are cool, especially on the hogback, western red moist, and somewhat more finely tex- 28 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tured than that of the pine-clad uplands. ula), little bluestem {Anclropogon sco- These habitats support American elm, parius), big bluestem (A. gerardi), cottonwoods (Poftiliis cleltoides), box- needle and thread grass {Stipa comata), elder (Acer negundo), willows (Salix porcupine grass (S. spartea), buffalo sp.), serviceberry (AmelancJiier aJnifo- grass {Biichloe dactyloides), Indian grass lia), red-berried elder {?>amhucus pu- {Sorga.struni avcnacciim), and Kentucky hens), blue flag [Iris missoiiriensis), bluegrass. Forbs include pricklypear Solomon's seal {Fohjgonaium hijlorum), cactus, pin-cushion cactus, soapweed carices {Carex sp.), and rushes (Junciis (Yucca glatica), sweet clover {Melilotus sp.). officinalis), blue vervain {Vervena has- Mountain Prairie Association.—On a tata), lamb's-quarters {CJienopodium al-

bum , sunflowers Helianthus hare- few rolling uplands at fairly high ele\'a- ) ( sp. ) , bell tions, an isolated prairie type of habitat (Campanula rotundifolia), western develops in areas where soil texture, salsify (Trogonpogon dubius), prairie moisture, and depth reach a favorable sandreed (Calamovilfa longifolia), balance. Examples of these grasslands skunkweed (Croton texensis), scarlet and wild alfalfa are the Gillette and Reynolds prairies, gaura (Gaura coccinea), and the Bald Hills (Fig. 11). Green (Psoralea tenuifiora) . Dominant browse are su- needlegrass (Stipa viridiila), western species ehokecherry, skunkbush ndae wheatgrass (Agropyroti sinithii), blue (Rhus trilobata), wormwood sage- brush dracuncidus A. grama (Boufelonra gracilis), timothy (Artemisia and A. (Pldetim prafense), red-top grass (Ag- dracunuloides) , fringe sagebrush ( rostis palustris) and brome grass {Bro- frigida), mountain mahogany (Cerco- western mus sp.) occur with silver sagebrush carpus montanus), sandcherry {Artemisia cona), brittle pricklypear cac- (Prunus besseyi), and wild rose (Rosa Scientific of in tus (Opiintia fragilis), common prickly- ivoodsii). names plants cactus O. the resume follow the of Wetmore pear ( compressa) , plains prick- usage cactus and and lypear (O. pohjcantha) , pincush- (1968) Pettingill Whitney ion cactus ( 1965 ) , \\'hereas vernacular names {Coryphantha missouriensis) , gen- follow the of Over goldcnrods (Solidago sp.), Indian paint- erally usage (1932). brush (Castilleja sidpluirea), mariposa Professor Ronald L. McGregor and asso- ciates at The Universitv of Kansas cur- lily [CaJchortus nuttaUii), gland stem are the flora of the (Adenocaidon hicoJor), and asters {Aster rently studying Black Hills sp.). region, and I am grateful to them Great Plains Prairie Association.— for \erifying the nomenclature or sug- for use here. Many tongues of the surrounding plains gesting appropriate changes extend into the Hills, especially from the south. The prairie-forest border is be- PLEISTOCENE HISTORY tween 3500 and 4000 feet in elevation. Pre-Wisconsin Events.—The Pleisto- Graves (1S99) indicated that in areas cene Epoch was characterized by a series where trees occur in xalleys and on of climatic fluctuations throughout the north-facing slopes, the prairie-forest world. (At least four southern cool plu- border is natural. However, at localities vial periods occurred conciurently \\'ith where trees occur on ridges instead of northerly continental glaciation. Subse- valleys, the prairie-forest border prob- quent to each of these episodes, a com- ably is due to fire. Prairie grasslands paratively warm, dry interglacial inter\'al formerly pre\'ailed in the Red Valley. followed.) As a consequence of these Vegetation of the prairie is far from distribution and uniform in its composition. Dominants oscillations, speciation of elements of the boreal among the grasses are western wheat- many and tem- grass, green needlegrass, blue grama, perate biotas were markedly affected. sideoats grama {Bouteloura ciirtipend- Prior to the Pleistocene, mountain- TURNER: MAMMALS OF THE BLACK HILLS 29 building in the Rock\' Mountain region the Missouri River, roughly paralleling probably created a rain shadow in the the river channel (Flint, 1957). Black Hills Further region. uplift during Apparently, these glacial periods the Pleistocene caused entrenchment of were characterized by cool, moist condi- streams around the edge of the Hills tions that resulted in the southward dis- (Darton, 1909). Previously, the Chey- placement of a northern biota. Fossil enne Ri\er and other streams to the remains of typical Hudsonian and Ca- south of the Hills flowed to- probably nadian zone mammals (including various the Gulf of Mexico. the ward Thus, shrews, microtines, sciurids, caribou, of the Missouri Ri\ er had present course muskoxen, moose, and the marten) have not been attained yet (Flint, 1955; been described from Wisconsin deposits

Lemke et 1965 . al., ) far to the south of their present distri- Because of relatively low elevation butions (Banfield, 1962; Barbour, 1931; and low precipitation, montane glaciers Benninghoff and Hibbard, 1961; Cush- did not form on the Black Hills (Hay- ing, 1945; Findley, 1953; Guildav, 1967; ward, 192S; Mcintosh, 1931). Nonethe- Hay, 1923, 1924; Hibbard, 1949, 1970; less, Darton (1906) and others reported Hibbard and Hinds, I960; Jakway, 1958; that Cordilleran glaciers formed in the Murray, 1957; Schultz, 1934; Schultz et Bighorn Mountains between 9500 and al., 1951; Semken et ciL, 1964; Skinner, 11,000 feet and flowed down to 6500 1942; Stearns, 1942; Wilson, 1967). There feet. There is no evidence that conti- is good evidence that during a part of nental entered each glaciers Wyoming ( Long, interglacial period (Aftonian, Yar- 1965), but at its maximum, about 18,000 mouth, and Sangamon), subtropical cli- BP, the Wisconsin ice sheet terminated mates extended farther northward than

miles east of the Hills . tran- about 150 Black now ( Hibbard, 1960 ) Subsequent (see below), and that part of South sition to subhumid, mesothermal climate Dakota west of the Missouri River es- led to each following glacial episode.

direct action 1957 . Rule states that within caped glacial ( Flint, ) Bergmann's Intense frost evidently occurred within a given species of warm-blooded animal, 50 to 100 miles of the ice sheet, making body size increases with latitude, thus the Hills region subject to periglacial in- implying an adaptation that presumably fluences. Erosion, deepening of stream serves to decrease the surface to volume channels, deposition of sands and gravels, ratio, thus conserving body heat in cold and development of soils also occurred environments. Guilday et al. (1964), in that region during the late Pleistocene. Flibbard (1963), and Parmalee (1967) evidence of shifts in The Nebraskan and Kansan glacia- presented popula- tions covered most of South Dakota east tions in some late Pleistocene faunas in climatic of the Coteau du Missouri, whereas the response to change; thus, larger northern taxa were smaller Illinoian glacier reached only the ex- replaced by southern relatives coincident with ame- treme southeastern corner of the state, lioration of cool climatic con- near Hartford (Lemke et al, 1965). This post-glacial ditions. Modern of these record is based principally on till found representatives same a similar of overlying presumed Pearlette Volcanic species display pattern Ash of late Kansas or Yarmouthian age variation. (Schultz and Smith, 1965). Distribution Although details are conjectural, evi- of Pre-Wisconsin till indicates that gla- dence of the displacement of biotic as- ciers advanced chiefly via the James semblages during the Pleistocene seems River and Red River lowlands. The Wis- impeccable. Voss (1939) estimated that consin ice front, as evidenced by accumu- the biota inhabiting four million square lation of drift deposits, extended from miles was either obliterated or displaced southeastern South Dakota, northwest- during this period. Amelioration of cli- ward toward North Dakota, just west of mate during the interglacial stages al- 30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

lowed migration northward; reversal of (Bryson et ah, 1970); thus, winters were climatic trends resulted warmer than at Absence or during glaciation present. ( only in displacement southward. Some iso- weak development) of hot dry wester- lated relict colonies were left in favor- lies, which did not begin to increase in able or tolerable locations along the strength and effect until the beginning various routes of dislocation of the Boreal Period (MacGini- ( Bryson and Wend- tie, 1959). land, 1967), resulted in cooler, more moist summers than at Massive shifts of biotic assemblages present. In pos- occurred several times in response to tulating climates and life-zones coinci- dent with Wisconsin changing climates and resultant glacia- glaciation, Dillon tion. Although the sequence of events {loc. cit.) proposed a depression (from the of 10-25 F in mean annual for each glacial episode and subsequent present) for the interglacial stage presumably was simi- temperature unglaciated portion of the Northern Great Plains. His lar, it is necessary to consider only the pro- is based on the that effects of the last continental glacier and posal assumption 45^ was the mean ensuing post-glacial events (including maximum temperature under which a could continue additional minor oscillations) to explain glacier the composition of the Recent biota on growth. Because of their higher eleva- the Black Hills the Black Hills. Effects of temporary tion, currently are cooler in warmer in and re- isolation imposed by Pre-Wisconsin gla- summer, winter, ciations would have been damped by ceive greater annual rainfall than do the chmatic reinvading elements through genetic surrounding plains. Therefore, conditions swamping that would be permitted by suggested for the plains dur- the next advance and retreat. Much of ing Wisconsin time may be somewhat the biota on the Hills is endemic at the extreme for the Hills, which presumably subspecies and varietal levels of diver- would have been more mesic and less gence. Had the current biota been effec- subject to wide seasonal fluctuations in than other re- tively and totally isolated by Pre-Wiscon- temperature periglacial sin episodes, endemism at the species gions. has varied level presumably would be more pre- Speculation concerning the nature of the dominant in the Black Hills. In addition, vegetation immediately the ice sheet. Tundra the warm hiatus of each interglacial bordering (Mar- stage seemingly would have affected the tin, 1959; Watts, 1967), grassland boreal and montane components ad- (Shimek, 1948; Kendeigh, 1961; Frey, 1965 and forest Clements and versely, causing extensive extirpation and ) ( Chaney, have been eliminating potential centers of dispersal. 1937; Flint, 1957; Byers, 1961) assigned to the of the ice Wiscomin Events.—The precise cli- periphery front. matic conditions in South Dakota and Rudd that the con- Wyoming during maximal continental ( 1951) postulated tinental were too cold and to and Cordilleran glaciations remain open plains dry continuous to question. Dillon (1956:168) proposed support forest during glacia- tion. The forested that dry, cold expanding air masses de- margins of present- scended off the icecap as a result of anti- day montane glaciers and the previously cyclonic circulation. This resulted in a mentioned evidence for warmer winters mean annual rainfall of about 10 inches and more mesic summers on the plains maximal in the periglacial regions. However, the during glaciation contraindicate pollen spectrum provides evidence of a Rudd's proposal. somewhat more mesic environment and on Potzger Tharp ( 1947) and Deevy the Northern Great Plains maxi- 1949 first during ( ) emphasized the biogeo- mal glaciation. The northern ice sheet graphical implications of spruce pollen was an impediment to the southward occurring far south of its present distri- flow of cold Arctic air onto the plains bution during glaciation. Additional TURNER: MAMMALS OF TliK BLACK HILLS 31 palynological studies have substantiated ous forest in the east or pine savanna these earlier reports, and confirmed that and mixed gallery forest in the west, and a belt of fir {Abies sp.), hemlock {Tsuga finally to steppe or pine savanna farther south. sp. ), and spruce {Picea sp.) existed across the central portion of the United Pollen diagrams from artesian-spring States during maximum glaciation. Birch marshes near Muscotah, Kansas, and {Betula sp.), alder (Alnus sp.), and from southern Minnesota provide no evi- Laiix also. dence of tundra the ice sheet tamarack ( sp. ) were present bordering Sedges (Cypemceae) and sagebrush during the Full-glacial Period (Wright, {Artemisia sp.) were the main herb con- 1970). However, pollen and plant mac- stituents of the boreal forest (Wright, rofossils indicate a narrow zone of 1970). Throughout the southern states, tundra did exist between the boreal for- from northern Florida to Texas, boreal est and glacial front in northeastern plant species were present but repre- Minnesota at a slightly later time (Watts, sented a more limited percentage of the 1967). total Pleistocene pollen-rain. For ex- The boreal flora evidently followed ample, Hafsten (1964) noted that pres- the glacier as it retreated northward in ent-day grassland areas of west Texas response to climatic change. Deteriorat- were occupied by pine during late Wis- ing spruce forest and the succeeding consin time. Recent studies (Watts and vegetation on the Great Plains varied Wright, 1966; Kapp, 1970; Wright, 1970) from east to west corresponding to suggested that in the Northern Great changes in moisture and from north to Plains, at least the Dakotas and portions south in relation to temperature gra- of Kansas and Nebraska were occupied dients and shifting isotherms. by a boreal spruce forest in Wisconsin The boreomontane character of the time. Fossil spruce wood in glacial drift Black Hills was established during the from Brookings County, South Dakota, Late-glacial times. This interval, which has radiocarbon dates greater than 30,000 followed maximal advance of the Wis- et BP (Lemke al, 1965:21). The current consin glacial front, was a time of stag- latitudinal altitudinal limits and of spruce nation, retreat, and periodic minor re- distribution approximate the 70 F July advance of the ice sheet. Presumably average isotherm (Kapp, 1970). there was a general withdrawal of boreo- Braun (1951; 1955), Kendeigh montane elements from the Northern (1961), and Thomas (1951) postulated Great Plains, accompanied by an increas- a temperate biota inhabiting the margins ing invasion of temperate species. Dis- of the glacier in the Great Lakes region junct populations of boreomontane mam- and eastern North America, and argued mals were left behind the retreating ice for relatively little vegetational change border in the Black Hills and in pockets south of the ice cap. In contrast. Gush- of suitable sites across the plains. ing (1965) and Gurtis (1959) favored Coincident with the northeastward an azonal mixed coniferous-deciduous retreat of the continental ice sheet, the vegetational margin south of the ice boreal spruce forest may have spread front. On the bases of pollen stratig- over those parts of the Northern Great raph\ , periglacial geomorphology, plant Plains that previously might have been macrofossils, and meteorological princi- treeless. The impressive cover of paleo- other authors ples, (Dillon, 1956; Mac- sols, sand dunes and loess (Peorian and 1959 else- Ginitie, 1959; Martin, and Bignell) deposited over much of the where; Guilday et al., 1964; Bryson et region in late Pleistocene time opposes zones of al., 1970) proposed vegetational the hypothesis of a dense periglacial transition. These transitional zones pro- forest, at least northward from western gressed southward from apparently tree- Nebraska to the glacial border (Smith, less tundra to boreal woodland, decidu- 1965; Wright, 1970). Ecogeographic dis- 32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

tiibutional patterns of steppe-related Sand Hills, eventually reaching the Black mammals also refute the presence of a Hills. dense woodland throughout the plains Dillon (1956:173) mapped the hypo- at this time (Hoffmann and Jones, 1970). thetical distribution of white spruce dur- Schultz et at. (1951:28) suggested that ing Wisconsin time and suggested that the formation of loess indicated a time boreal elements entered the Hills via a of "reduced vegetative cover." In north- link with the Rocky Mountains. How- eastern Kansas, the nature of the loess, ever, were this the case, the species in the molluscan fauna of the loess, and the Hills probably would be Engelmann's the pollen diagrams from Muscotah indi- spruce {Picea engelmonni) (Halliday cate wooded environs. Farther west, and Brown, 1943). Thus, the white the loess and associated molluscan fauna spruce in the Black Hills seemingly suggest a prairie environment interlaced was displaced originally from the Cana- with gallery forests that provided habi- dian or Hudsonian life-zones to the north. tats Evidence of coniferous for nongrassland species ( Leonard species through- and Frye, 1954). Kendeigh (1961) noted out South Dakota during some part of that the nature of the loess of the plains the late Pleistocene implies that boreal was indicative of grassland abutting di- components could have invaded the Hills rectly against the glacial front. How- from several directions, following cli- ever, meteorological evidence suggests matic change and resultant glacial re- that grassland \egetation in the Northern treat, rather than solely from the west. Great Plains during the Late-glacial in- The present boreal elements in the Hills terval may have been restricted to the are disjunct from contiguous boreal for- more arid rain shadows of the Rocky est, which is located some 435 miles to Mountains, in western Kansas and east- the north (Buttrick, 1914). ern Colorado (Bryson and Wendland, The cordilleran-montane biota also 1967). entered the Black Hills during the Full- Sand dunes are not well correlated glacial Period. Regional snow lines and with Pleistocene events, but the Bignell montane biotic zones were depressed Loess is radiocarbon dated at about vertically about 4000-4500 feet during 12,600 BP (Watts and Wright, 1966), that time (Martin, 1959:394; Richmond, and maximum loess deposition occurred 1965:228; Webb, 1965:457). Richmond in Illinois and Iowa from 20,000 to {loc. cit.) suggested a decrease of 16- 14,000 BP (Wright, 1970). Smith (1965) 17 "F in summer temperature concurrent suggested that large transverse sand with the displacement. Coincident with dunes were formed by periglacial winds these events, the cordilleran-montane ele- from the north, at a time when unfor- ments extended considerably downslope, ested terrain could not interfere with eastward and southward, in\'ading the dune development. Juxtaposition of the Black Hills, Laramie Mountains, and vast area of desertlikc dunes in northern Great Plains along a wide front. Ensu- Nebraska in contrast to the well docu- ing climatic changes during Late-glacial mented occurrence of a boreal spruce and Holocene times caused retreat of forest in other regions of the plains dur- these components toward their previous ing Late-glacial time is compromised by centers, leaving relics in the Hills and Wright (1970). Retreat of the ice front, other montane environs, on escarpments, with subsequent decrease in velocity of and in favorable mesic sites on the plains. periglacial winds, permitted the en- Montane floral elements of the Black croachment of vegetation that stabilized Hills currently are disjunct from those to the dunes. Presence of the subalpine the west, the nearest populations residing spruce forest in the Black Hills may indi- in the Big Horn Mountains of Wyoming, cate that boreal vegetation then spread about 150 miles distant. rapidly over (or around) the Nebraska Subsequent climatic oscillations were TURNER: xMAMMALS OF THE BLACK HILLS 33 not of equal direction or intensity. The concerning biotic events follow Hoff- chronology of climatic events depicted mann and Jones (1970), Watts and in Table 2 is based on episodes defined Wright (1966), Wells (1970a, 1970b, by Reid A. Br^son and others (Biyson 1970c), and Wright (1968, 1970). and Wendland, 1967; Baerris and Bry- Post-Wisconsin Events.—Climatic et son, 1965; Bryson al, 1970). More change that finally terminated the Pleis- recent episodes ha\'e been further clari- tocene epoch brought about rapid phyto- fied studies of cultural by history (Leh- geographic change. On the eastern por- of historical mer, 1970) and droughts tion of the plains, spruce forest suc- (Tomanek and Hulett, 1970). Notations cumbed to a brief increase of alder and

Table 2.—Chronological events since maximal Wisconsin glaciation.

PLEISTOCENE EPOCH

Full-glacial Period (30,000 to 13,000 BP) —Maximum advance of Wisconsin ice sheet; sum- mers moister and cooler, and winters warmer than present. Boreal spruce forest over much of Northern Great Plains; north-central Nebraska to glacial border presumably treeless; members of present plains biota occupy steppe and savannah conditions to south. Late-glacial Period (13,000 to 10,500 BP) —Stagnation, retreat, and periodic minor readvance of ice sheet; culmination with Valders Readvance about 10,600 BP; summers moister and cooler, and winters warmer than present. Boreal forest still present; grassland restricted to arid rain shadow of Rocky Mountains; complex of steppe, taiga, and timdra biota in western section; pine sa\annah and steppe still south of Northern Great Plains. HOLOCENE EPOCH

Pre-Boreal Period (10,500 to 9650 BP) —Shift in atmospheric circulation; retreat of ice sheet; low corridor opening l)etween Arctic and Great Plains; frequent and intense polar storms; westerlies de\'eloping; increasingly continental climate with summers drier and warmer, but winters colder than present. Replacement of boreal forest by steppe on plains, and by deciduous elements to east. Boreal Period (9650 to 8450 BP) —Increased frequency and strength of westerlies and polar storms; continental climate with summers drier and warmer, but winters colder than present. Spread of grassland northwestward south of ice front. Atlantic Period (8450 to 4680 BP) —Increased southward flow of Arctic air; westerlies attain maximum strength; summers arid and warm, with increase in moisture late in period, but winters colder than present. Maximal eastward penetration of grassland; boreal and deciduous forest north of present limits. Sub-Boreal Period (4680 to 2890 BP) —Southward shift of Arctic frontal zone; chinooks less frequent east of Rockies; temperatures cooler; climate similar to present by end of period. Steppes retreat to present limits; southward shift of boreal biota; lower tree line descent down eastern slopes of cordillera. Sub-Atlantic Period (2890 to 1690 BP) —Shift in upper-air anticyclonic circulation; influx of tropical air resulting in moister climate than present. Possible influences on biota north- east\\'ard from Great Basin. Scandic Period (1690 to 1100 BP) —Transition period back toward early Atlantic conditions, becoming warmer and drier once more. Neo-Atlantic Period (1100 to 760 BP) —Sidatropical anticyclones cause influx of moist tropical air; warmer climate continues. Conditions favorable for corn agriculture; horticultural villages established in Missouri River Valley. Pacific Period (760 to 410 BP) —Increased westerlies across North America; resultant drier climate (30 to 50 percent decrease in summer precipitation); return to Neo-Atlantic conditions later in period. Decreased extent of occupation of Missouri River Valley by village tribes; return of occupancy later in period. Neo-Boreal Period (410 to 115 BP) —Westerlies and polar storm tracks shift southward and intensify; summers drier and cooler; increased moisture later in period. Small temporary villages with marginal economy in Missouri River Valley; larger more permanent villages later in period; effects of European man after 290 BP. Recent Period (115 to present) —Increased westerlies resulting in warm, semiarid climate at present; short periods of drought commencing about 102, 91, 60, and 18 BP. 34 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

birch, which in turn were replaced by ponderosa pine with lower synusia of other temperate deciduous elements such semidesert slnubs and grasses. as Ulmus hazel The available evidence a elm ( sp. ) , hop hornbeam, suggests {Conjlus sp.), and oak {Quercus sp.), more or less azonal plant and animal with parklike openings inhabited by community similar to that postulated by herbs and grasses (Watts and Bright, Jones (1964:23) for Nebraska during 1968; Wright, 1970). To the west in maximal glaciation in the Northern Great Kansas, Nebraska, and the Dakotas, the Plains during Late-glacial and early post- boreal forest gave way diiectly to prairie glacial times. Both prairie and forest species (Wright, 1968, 1970). Two rec- components may have occurred in the ords of tundra-related mammals noted same general area to form a savanna-Hke for South Dakota suggest an environment landscape. The relative frequency of equivalent to present day Hudsonian and boreal elements decreased from north to Canadian life-zones. Martin (1959:398) south and deciduous elements decreased reported a muskox (questionably Ovi- from east to west. Relatively cold-toler- hos) with a radiocarbon date of 9700± ant, widely distributed temperate species 600 BP, and Green and Lilligraven of grassland affinity may have been seg- (1965:48) Hsted a fossil caribou from the regated ecologically in accordance with late Pleistocene in Gregory County. Re- local conditions (Jones, loc. cit.). The lict populations of boreal and montane supposition of an admixture of northern species of plants and animals on the and southern elements in the plains re- Black Hills also attest to these environ- gion also is in agreement with the view- mental conditions (Jones, 1964:21). point of Smith (1957:207). Time of demise of the boreal spruce During warm, relatively moist inter- forest in the plains region has been re- vals of post-glacial time, such as in the corded by various authors, as follows: Sub-Atlantic or Neo-Atlantic periods, Muscotah, in northwestern Kansas, elements of the eastern deciduous forest 15,880-11,340 BP (WMght, 1970); Rose- and tall-grass prairie biotas probably bud, in the Sand Hills near the Nebraska- extended westward far onto the plains state BP via South Dakota hne, 12,600 ( eastern deciduous species mesic gal- (Watts and Wright, 1966); Rosebud in lery forests along river systems and val- western Kansas and Nebraska, 11,400- leys, and tall-grass components on the 9100 BP (Ruhe, 1970); Pickerel Lake, uplands). Hardier boreal elements re- in northeastern South Dakota, 10,670 BP maining from the Late-glacial Period (Watts and Bright, 1968); Pickerel Lake presumably could have survived these in Iowa, 8000-7000 BP (Ruhe, 1970). somewhat mild climatic conditions in The tree-prairie transition is dated by comparatively undisturbed, disjunct pop- at in the south- ulations. the Wright ( 1970) 12,000 BP Jones (1964) interpreted ern portion of the Great Plains and 9500 relatively large amount of fossil pine BP in the northern section. pollen described from the Nebraska Sand Hills Sears to indicate that Later occurrence ( 5040 BP ) of wood- by (1961) boreal survived lands (with somewhat more pine than species early post-glacial time only to be excluded by ensuing mid- present today) on the plains of north- post-glacial periods of maximum warmth central Nebraska and South Dakota is and dryness. The eastern deciduous bi- well documented also (Kapp, 1970; ota may have mingled with residual cor- Sears, 1961; Watts and 1966; Wright, dilleran elements (from Pleistocene or Wells recorded Wright, 1970). (1970b) Sub-Boreal times when the biota spread the flora of the Laramie mid-post-glacial eastward and southward down the slopes Basin, southwest of the Black Hills, as of the Cordillera) at points throughout being an open xerophilous woodland the Great Plains (Webb, 1965). dominated by western red cedar and Conversely, arid post-glacial periods TURNER: MAMMALS OF THE BLACK HILLS 35 were a time of extirpation o£ the boreal drought environment promoted an east- and eordilleran-montane elements that ward expansion of the steppes. In time, survived from late Pleistocene time on amelioration of dry conditions resulted the Northern Plains; only the most tol- in retreat of grasslands to their present erant species survived in small, isolated limits. This sequential series of events populations. Much of the unique biota can be traced in pollen diagrams of Pick- of the Black Hills may have been re- erel Lake and Muscotah (Watts and stricted during these periods. Outlying Bright, 1968; Wright, 1970). Wells representatives probably could not sur- (1970b) described the present flora of vive the effects of the hot, dry climate; the Laramie Basin as desertic in physiog- the milder, more stable climate in the nomy with strong resemblance to winter- Black Hills presumably afforded a suit- cold deserts of the Great Basin and Colo- able refugium for these elements. Those rado Plateau. Change from a xerophi- species of eastern and southern affinities lous woodland to a semidesert shrubland that had spread into the Northern Great in post-glacial time denotes a secular Plains area during the humid climatic trend toward an increasingly arid climate episodes either found suitable refugia east of the Laramie Front Range. Re- during the time of exapotranspirative cession of the boreal forest barrier that stress, or were subsequently excluded. existed between the Great Basin and During the mid-post-glacial time of Wyoming Basin during the Wisconsin maximal xeric Hoffmann and 1970 aridity, plant species sup- ( Jones, ) opened ac- planted mesophytic vegetation; these cess for cold-desert species to the North- species fragmented formerly continuous ern Great Plains. Shifts in upper-air areas of deciduous vegetation and per- anticyclonic circulation northeastward mitted encroachment of arid-tolerant from the Great Basin in the Sub-Atlantic trees and nonarboreal plants (Smith, and resultant influx of tropical air also 1965). Gleason (1923) invoked an east- may have influenced the biota of the ward extension of the prairie to explain region. For example, evidence of exten- the relict pattern of prairie plant distri- sive introgression between the bur oak in in the Black Hills and Gambel's bution (prairie peninsula) the Mid- oak ( Qiier- west, and postulated the extension prior cus gambelii) to the southwest, in the to the invasion of deciduous species in central Rocky Mountains, presupposes a post-glacial time. Transeau (1935) in- massive northeastward migration of the dicated that the extension was subse- cold-intolerant Gambel's oak during a

to initial warm moist 1970b . quent the invasion of the decid- period ( Wells, ) Such uous forest in mid-post-glacial time. conditions were not current during east- Available evidence implies that maximal ward and southward displacements of eastward penetration of the grassland eordilleran-montane elements. occurred in the Atlantic Period. How- The sequence of episodes in the his- ever, as with the north-south glacial dis- tory of native cultures of the Missouri placement of the biota, the west to east River Valley in the Dakotas (Lehmer, dispersal of prairie species may have 1970) and the Mill Creek culture of Iowa been several times. repeated (Bryson et al., 1970) implies a close cor- Thus, during milder intervals of post- relation between climatic and cultural glacial time, species having eastern and changes in near-modern time. Modera- possibly southern affinities spread onto tion, culminating in the present climatic the Northern Great Plains and Black conditions, has been interrupted by a Hills, and mixed with residual boreal gradual warming trend on the North and eordilleran-montane constituents. American continent in the last 100 years During subsequent arid climatic epi- (Dorf, 1959, and others). Periodic, ex- sodes, these immigrants either inhabited tended droughts have resulted in changes suitable refugia or were eliminated when of the grasslands and associated fauna 36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

(Tomanek and Hulett, 1970). Wooster southeast of the glacial front or possibly (1935, 1939) and Gier (1967) noted that as mosaic distributions in a prairie-forest some mammals, such as the prairie vole savanna. When one realizes that most almost have a of (Microtiis ochrogaster) , disap- grassland species major part pear in some areas under arid regimes, their diverse ranges as synusial compo- whereas others, such as the deer mouse nents of serai stages of forested en\'irons, seem rela- the of the biome be- (Peromijscus maniculatus) , uniqueness prairie tively unaffected by extended drought. gins to fade; consequently, the grasslands The Great Plains of Recent time is should be regarded as consisting of a an extensive grassland on deep, trans- derivative biota. ported soils, corresponding to flat or Climate alone does not account for a gently rolling topography. The gram- grassland climax vegetation. In areas inoid flora is defined by a predominance where grasslands were planted in Ne- of a few widespread species and a pau- braska with upland tree species native city of endemics, virtually none of which to scarp woodlands, rather extensive for- are grasses (Wells, 1970c). The associ- est became established (Wells, 1965). ated insect fauna (e.g., range grasshop- Furthermore, Potter and Green (1964: pers and lataline grass-feeding leafhop- 22) found evidence of ponderosa pine, pers), has similar distributional charac- as seedlings and young saplings, invad- teristics; however, the fauna has a much ing the grasslands in North Dakota. A greater species diversity in certain serai similar invasion can be observed on the stages of surrounding forested biomes prairie of Wind Cave National Park, than in comparable grassland communi- where grazing by bison checks the re- ties (Ross, 1970). Generally, trees are re- production of ponderosa pine. Thus, in stricted to riparian habitats as gallery at least certain locations, the plains are forests along water courses; in addition, quite capable of supporting nonriparian nonriparian woodlands occur on escarp- woodlands (at least of xerophytic coni- ments and other topographic breaks fers). Physiography may be as impor- throughout the grassland province of tant a factor as climate in affecting the central North America. distribution of extensive, treeless grass- Clements and Chancy (1937) indi- lands. Stewart 1951 noted that theoretical cated that the prairie species probably ( ) climatic climax in savanna developed after Miocene time in the vegetation and areas central parts of the continental United grassland probably postdates States due to drier conditions created by the arrival of man. The Tule Spring, Nevada dated at mountain building to the west. How- site, +23,000 BP, indicates that was ever, Wells (1965, 1970a, 1970b, 1970c) clearly prehistoric man several present maximum glaciation in and Ross ( 1970 ) presented argu- during the Wisconsin The fre- ments for relatively recent tenure of the (Martin, 195S). of which is as- central grasslands. Accumulated evi- quency fires, generally dence indicates that these extensi\'e tree- sumed to have increased with the arri\ al of is well documented in his- less steppes probably have not had a con- man, early torical literature. and tinuous existence since the mid-Tertiary. Scarps abrupt breaks have ser\ed as The abundant, nonarboreal pollen re- topographic may from fires for the nonri- ported by Kapp (1970) from interglacial refugia grass sediments of the Sangamon interval may parian \\'oodlands of the plains region be from restricted refugia; the occur- (Wells, 1965, 1970a, 1970b, 1970c). rence of the pollen does not necessarily Forest fires may ha\'e terminated imply the presence of widely distributed broad expanses of boreal forest that steppe vegetation. During maximal Wis- were already suffering evapotranspira- consin glaciation, grassland species sur- tive stress from the effects of warm post- vived in suitable refugia southwest and glacial climate. These newly opened TURNER: MAMMALS OF THE BLACK HILLS 37 — expanses subsequently were invaded by Table 3. Distril:)utional patterns of the flora of the Black Hills, as noted liy Hayvvard ( 1928) a biota ha\'ing strong ecological affinities andMcIntoch (1931). with nearby forested biomes. Interaction of topography, climate, and subsequent Biographic origin Percent affinity prairie fires apparent!)' has resulted in Western 25-30 the present ph\ siognonn' of the Northern Widespread 22-34 Great Plains. See the several of papers Great Plains 17-26 Wells cited above for additional argu- Eastern 9 ments concerning these points. Northern 6 The introduction and development of Southern 4—5 modern ci\ilization and agriculture in the no doubt ha\'e resulted plains region Table 4. —Distributional patterns of lichens of in extirpation of several species, endan- the Black Hills as noted by Wetmore (1968). gered others, and caused widespread, drastic modification of habitats that frag- Biogeographic origin Percent affinity mented the of other ranges yet species Arctic-Boreal 44.4 (Smith, 1965). In addition, man has in- Pan Boreal 24.9 troduced many exotics (e.g., Kentucky Pan Temperate 5.4 Western 4.9 bluegrass) and thereby possibly caused Temperate Arid Southwestern 4.4 the exclusion of native or shifts species Pan North America -^ 3.9 in their respective ranges. Southern Rockies- The Black Hills flora is diverse, with Alleghenian-Great Lakes 3.9 Eastern inany species occupying a small geo- Temperate 3.4 Eastern Boreal 2.9 graphical area. For example, Froiland Grassland 1.5 identified 20 kinds of (1962) willows Western Boreal 0.5 (Salix sp.) of diverse geographic origin in the Hills region. This diversity is due partly to the varied topography and hab- Examples of boreal species are paper itats available, and partly to the past birch, twin-flower, Venus' slipper, bunch- movement of great floral assemblages berry, squashberry, and white spruce. during late Tertiary and early Pleistocene Montane species include Oregon grape, times (Wetmore, 1968). However, the Indian paint-brush, mariposa lily, skunk- Black Hills are not unique in having dis- bush sumac, buffaloberry, white spruce tributional limits of so many northern, and ponderosa pine. Fernald (1935) southern, eastern and western species. suggested that a few western species— The for stem and probable geographic origin of — example, gland huckleberry the vegetative components of the Black may represent pre-Pleistocene floral Hills are listed in Tables 3 and 4. The elements. presence of many widespread species Great Plains components (17-26%) (22-34%) can be attributed to the fact are to be expected because of the loca- that plants with wide ranges of tolerance tion of the Black Hills, which are entirely overlap distributionally within the Hills surrounded by plains. These plants are region. Most northern or boreal (6%) found in the foothills, in the drier mead- and western or montane (25-30%) spe- ows, and open parklands. Examples are cies were displaced southward or east- blue grama, Indian grass, porcupine ward by advance of continental and grass, and western sandcherry. Eastern Cordilleran ice sheets, respectively. species (9%) presumably reached the These species became stranded on the Hills during the Sub-Atlantic or Neo- Hills by migrating up the slopes after Atlantic intervals via gallery forests along glacial retreat; although many species water courses and uplands. These may presumably were extirpated in the At- be Ozarkian and Alleghenian elements of lantic Period, some remain currently. the eastern deciduous forest (Braun, 38 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

1947). Examples are hop hornbeam, americanus), and prairie dog, have been American elm, boxelder, Solomon's seal, reduced drastically. When European green ash, and bur oak. man first moved into the Black Hills, The southern components (4-5%) big-game mammals, fur-bearing mam- probably arrived at about the same time mals, and smaller game mammals were as did the eastern elements. In part these abundant, but subsequently they have are derived from the old Madro-Tertiary become the object of exploitation. As flora that developed in the southwestern agricultural endeavors in the region in- United States in Miocene, Oligocene, creased, carnivores and rodents were and Pliocene times (Wetmore, 1968). poisoned indiscriminately. Fortunately, The driest areas of the foothills now are recent programs of wildlife management occupied by the following species: soap- and a more intelligent application of weed, brittle pricklypear cactus, skunk- rodent and predator control have less- weed, scarlet gaura, silver sagebrush, and ened the devastation by man. Estab- wormwood. The examples used in this lishment of Wind Cave National Park, resume were taken from Wetmore Custer State Park, and several other (1968). areas has served to protect and pre- The distiibutions of lichens in the serve remnants of the native fauna. Man Black Hills are vastly more complex than has reintroduced several species that for- the respective distributions of vascular merly were native to the Black Hills, plants because of the diversity of sub- such as the elk (Cervus canadensis), strates, exposed geological materials, ele- pronghorn, bison, and mountain sheep. vations and relief, climatic range, and In addition, the mountain goat, fox squir- vegetational types. The probable geo- rel (Schirus niger), house mouse {Mus of the lichens in the Black and rat nor- graphic origins muscuhis) , Norway (Rattus Hills are listed in Table 4. For a thor- vegicus), which are not native to the ough analysis of the complexities in- Hills, were introduced by man. Activi- volved in distributional patterns of lich- ties of man that have indirectly affected

. distribution of ens, see Wetmore ( 1968) the abundance and mam- malian species in the Black Hills are INFLUENCE OF MAN ON THE summarized below. For detailed infor- ENVIRONMENT mation concerning these actixities, the reader is directed to the "Black Hills The Black Hills region is one of the Area Resource Study," published jointly most densely populated areas of the by the Departments of Agriculture and Great Plains. In 1960, the Hills area con- the Interior in February 1967. tained an average of 6.2 persons per —The rush of 1874-79 square mile. Of the native population, Mining. gold to the in 57 percent were classified as urban, 30 gave impetus mining industry the Black Hills In the percent as nonfarm rural, and 13 percent region. early of and as farm rural. The presence of about 1900's, mining silver, tin, coal, crude oil flourished. The two World 6000 armed forces personnel and from four to five million annual tourists inten- Wars created a greater demand for vari- to sifies utilization of Black Hills environ- ous minerals and gave added impetus ments. the tungsten, sheet mica, feldspar, ben- and lithium industries in Changes wrought by man have been tonite, beryl, the Hills. The most recent boom detrimental to some mammalian species mining in with the of and advantageous to others. Large began 1951, discovery uranium ores in herbivores and carnivores, such as the Jurassic-Cretaceous bison and wolf Canis have sandstones. Production of nonmetallic ( lupus ) , been such as directly extirpated by man; populations commodities, sand, gravel, of other species such as the mountain crushed rock, cement, and clay, also are lion {Felis concolor), black bear {Ursus current industries of the region. Mining TURNER: MAMMALS OF THE BLACK HILLS 39 in\'aiiably alters the natural environment. moved into the Black Hills about 1900, For example, in processing bentonite, but the drought of 1910-11 shifted major the overburden is stripped off and the emphasis from general farming to the bentonite is turned oxer and stored in raising of lixcstock. Ranching became place for long periods of time. Excava- an important activity in the unforested tion of mining complexes and rock quar- uplands, foothills, and the Red Valley. ries also has altered the topography. Grazing now accounts for 75.1 percent Processing of gold in the Lead-Dead- of the principal land use in the Black wood area has resulted in pollution of Hills area; usually beef cattle and sheep man)' nearby streams. However, not all are marketed as feeder stock. alterations ha\'e been detrimental to the About 103,000 acres were under irri- regional fauna; for example, mines fre- gation in the Hills region in 1960. Crop- quently are utilized by woodrats and land comprises about 8.6 percent of the various species of bats. principal land use, with corn, sorghum, winter al- Timherinp,.—Cutting of timber com- wheat, spring wheat, barley, and the menced simultaneously with the Black falfa, hay being major dryland The farm in the Hills Hills gold rush, and at least two portable, crops. average area in 1960 was 2550 acres in steampowered sawmills existed in the extent, and sales of Hills by 1877. Intensive timbering caused agiicultural products amounted to more than 50 million dol- a rapid depreciation of the quality in lars. The en\ironment has been modi- native stands adjacent to mining opera- fied due to tions by 1897, when President Cleveland conspicuously agricultural created the Black Hills National Forest practices. The original prairie grasslands Reserve. Sale of federal timber was of the Red Valley have been altered by and the authorized by the Timber Reserve Bill, gi'azing upland grasslands (e.g., the and Gillette now enacted later in the same year. The Reynolds prairies) market and timber industry developed rapidly, sup- support hayfields, gardens, Wind Cave National Park is plying vast quantities of wood products pastures. one of the few areas wherein native to the mines, railroads, and other indus- large still exists in its tries of the region. The pulp and paper, prairie grassland nearly state. and the post and pole industries have original flourished recently. Forest production Fires.—The earliest records of fire in accounts for about 14.7 percent of the the Black Hills were about 1730 to 1740, principal land use in the Black Hills. when the entire area appears to have The total of commercial timber currently burned by a series of fires; in the period available in the Hills is estimated at five between 1790 and 1800, most of the Hills billion board feet; production in 1964 were burned again (Graves, 1899). was in excess of 21.8 million cubic feet, Lesser fires of considerable extent oc- contributing 6.5 million dollars to the curred in 1842, 1852, and 1875. From of for- regional economy. Exploitation 1880 through 1966, at least 70 fires in the ests is evidenced vast by lumbering by Black Hills burned more than 175,000 areas of scrubby second-growth timber acres of forest and grassland. Of these, and numerous open areas where burning 44 percent were known to have been has resulted from lumbering operations. caused by man and 18 percent by light- Removal of dead trees in clearing opera- ning, whereas 38 percent were of un- tions has eliminated potential den sites known cause of U. S. Forest for flying squirrels and red squirrels. (records Black Hills National However, populations of some mammals Service, Forest). in- the notable fires were those (e.g., deer and chipmunks) have Among more creased in areas of stumps, logs and trim- of Iron Creek in 1899 (38,000 acres), mings. Rochford in 1931 (21,640 acres), McVey Agriculture.—Homesteaders first in 1939 (21,875 acres), and Wildcat in 40 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

1960 (10,336 acres). Improved precau- decades has been the influx of four to tionary measures and modern fire-fight- five million tourists annually. These visi- ing techniques have prevented major fires tors have intruded into canyons, onto in recent years. Two smaller fires should the forested slopes, and upon much of be mentioned. Although the Deadwood the remainder of the relatively undis- fire burned only 4501 acres, it forced turbed en\'ironment in countless \\'ays. evacuation of that historic town in 1959. Recreation and tourism are the foremost The Headquarters fire in Wind Cave producers of revenue in the Black Hills; National Park in 1964 (4000 acres) more than 120 million dollars were con- greatly endangered the natixe wildlife tributed to the regional economy by this in the Park. A noticeable result of forest industry in 1966. People seeking recrea- fires is the opening of extensive new tion are attracted to the Black Hills by grassland, and the establishment of as- points of historic and geologic interest, pen and birch woodlands in burned by opportunities to hunt and fish, and by areas. Reforestation programs and nat- such scenic features as Custer State ural secondary regeneration have par- Park, Wind Cave National Park, Jewel tially revegetated the scarred hillsides Ca\'e National Monument, Devils Tower that resulted from past fires. National Monument, and Mount Rush- Recreation and Tourism.—Although more National Memorial. Demands for immeasurable in extent, perhaps the most recreation surely will continue to in- detrimental influence on the environ- crease in the future, as will exploitation ment of the Black Hills in the past few of the once pristine environment.

ACCOUNTS OF SPECIES

ORDER INSECTIVORA—Insectivores Central Basin and on the Limestone Family SORICIDAE—Shrews Plateau above 5500 feet. It is common locally in riparian associations and other In the Black the order Insecti- Hills, moist habitats throughout the Northern vora is one of the represented by species Great Plains. Except for three individ- genus Sorex. An additional species of uals ol:)tained in October and November, this occur in the and genus may Hills, all specimens examined were taken in another has been incorrectly reported warm months. from the area. In the Hills, this species is most nu- merous in montane habitats; it is locally Sorex cinereus Baird haydeni abundant in marshy areas, mossy bogs, Masked Shrew and other riparian associations of grasses, Sorex haydeni Baird, 1858, Mammals, in Re- sedges, rushes, aspen, birch, and \\ illow.

ports of explorations and surveys . . . from Many specimens were trapped near logs, the Ri\er to the Pacific Ocean Mississippi large rocks, or low shrubs located in . . ., 8 (1):29, 14 July (type locality re- dense grass along creeks such as Beaver, stricted to Fort Union, just west of conflu- ence of Missouri and Yellowstone rivers, Boxelder, Castle, Cold, Grizzly, Iron, Williams Co., North Dakota, by Merriam, Rapid, Rattlesnake, Spring, and Willow, N. Amer. 31 Farma, 10:60, December 1895). or in the xicinity of beaver dams. Spring- Sorex cinereus — haijdcni Jackson, 1925, Jour. fed bogs and other moist areas in aspen Mamm., 6:56, 9 February. and spruce woodlands also pro\'ide suit- Vernon Bailey (1888:436) first re- able habitat for these shrews. ported the masked shrew from the Black The availability of ground water un- Plills under the name Sorex personatus. doubtedly is one of the most important Known altitudinal range in the Hills ex- factors limiting the distribution of S. c. tends from 4500 to 6500 feet, but this haijdcni. A few indi\iduals ha\e been shrew is most widely distributed in the taken in unusual habitats; for example, TURNER: MAMMALS OF THE BLACK HILLS 41

a female was trapped on a rock ledge curs only in fully matured or old individ- near the top of a 30-foot cliff, and four uals in S. c. cinereus in Michigan. Three others were obtained on dry, rocky, pine- young females of age class two of Pruitt clad hillsides 200 feet loc. cit. approximately ( ) were obtained in summer and from the nearest source of water. Also, were gravid; these were taken on 25 June two shrews taken by Bailey (1888:436) (seven embryos), 29 June (five em-

were "at holes in the rocks on and 2 five . captured bryos) July ( embryos ) Pruitt top of one of the highest peaks." Popu- also indicated that the testes of reproduc- lation densities frequently are low in tively acti\'e males exceeded two milli- these marginal areas. Associated with meters in length; testicular length of an the masked shrew in moist habitats are immature male obtained on 3 July was 5. Zapus hudsonhis, Microttis longicoudns, While uncommon, such precocious repro- and M. penn.syhanicus. Feromijscus ductive activity has been reported in maniculatus, Chthrionoimjs gapperi, and other species of the genus Sorex. Young Eufaniia.s luinimtis have been taken with of the year were evident from late June these insecti\ores in drier environments. through mid-August. A member of a University of Kansas Due to the uniformity in season of collection to field party, J. R. Choate, buried 20 cans (mid-June early August), of the in in tall grass along Boxelder Creek, 20 most specimens examined were mi. W Nemo between 29 June and 5 summer pelage. However, a male ob- July 1967. The cans were buried 10 tained by Vernon Bailey near Sundance 13 paces apart with the openings at ground on August 1913 evidenced molt, with fresh winter level. A snap-trap was placed within five pelage covering about two- thirds of both and three feet of each can. These pitfalls (three dorsum venter; shrews captured) seemed to be a more others, taken by Merritt Cary in the Bear eflBcient method of obtaining S. c. haydeni Lodge Mountains on 22 June 1912, had than was snap-trapping (one shrew cap- just completed molt. tured); however, these data are insuffi- In considering geographic variation, cient and a more thorough application specimens of S. c. hoydeni from the of this technique in the Hills region is Black Hills were compared with individ- desiralile. Trapped insects may have uals from south-central Wyoming (Al- attracted the shrews to the pitfalls. bany and Carbon counties). North Da- Reproductive data are a\ailable for kota (Bowman and Walsh counties), S. c. haydeni in the Black Hills from mid- eastern South Dakota (Kingsbury, June through mid-July. Of 33 females Union, Marshall and Day counties), and northern captured during this period, 15 showed Nebraska (Sioux, Sheridan, Cra- no sign of reproductive activity; the same Cherry, and Keya Paha counties). was true for a female taken in Spearfish nial measurements were taken in the Canyon on 7 August. Mammae of a fe- manner described bv Jackson (1928:12) male obtained on 18 June were enlarged, and Findley (1955:5). Cranial breadths whereas another captured on 24 June of 29 females averaged somewhat greater 22 had an enlarged uterus, possibly indicat- (7.8±0.21) than those of males (7.6 ing recent implantation. Twelve females ±0.22). Otherwise, significant secondary taken in the period 25 June to 7 July sexual differences were not apparent nor within the Hills carried 5.2 (2-7) embiyos that were 5.9 was significant variation detectable. (2-11) in crov-'n-rump length. Four lac- population tating females were noted in the period Reflectance readings show that 27 June to 13 July. The testes of 14 males masked shrews vary geographically in captured early in the summer were 4.5 tone of color. Individuals from North (2-6) in length, whereas those of three Dakota and north-central Wyoming a\'er- taken earh' in August were 3. Pruitt aged paler than those from the Black

1954 : 36 that ( ) reported reproduction oc- Hills, wliereas specimens from south- 42 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY central Wyoming and Nebraska aver- sions, specimens from the Black Hills are aged darker than Hills specimens. intermediate between S. c. cinereus to Shrews from eastern South Dakota ap- the west in Wyoming and S. c. haydeni proached those from the Hills in colora- to the east in the Dakotas and Nebraska, but fall within the of \'ariation of tion ( Table 5 ) . No significant geograph- range ical difl^erences in hue were discernible. the eastern specimens and hence are S. c. Additional Both external and cranial dimensions assigned to haydeni. measurements listed in table of evince considerable geographic variation. (not 5) 67 adult shrews from the Black Hills Masked shrews from localities west of are: red reflectance, 54.3±0.52; the Black Hills have greater average percent percent green reflectance, 23.7 ±0.19; body, tail, maxillary tooth-row, and pal- percent blue reflectance, 21.3 ±0.32; hind ate lengths, as noted by Long (1965: foot 11.2±0.64; ear 6.4± 519). However, these individuals are length, length, 1.25; 3.9±0.94; condylobasal indistinguishable from eastern represent- weight, 15.5±0.28; cranial breadth, 7.7 atives on the basis of condylobasal length length, interorbital 2.8± and interorbital breadth. In addition, ±0.23; constriction, skull 4.7±0.33. the average rostral length is greater and 0.11; depth, the breadth across the maxillary proc- Specimens examined (136).—SOUTH DA- KOTA: Lawrence Counti/: Deadwood, 3 esses is significantly less than noted in (USNM); Dumont, 6100 ft, 3 (USNM); 2 mi from sites east of the Black specimens S Tinton, 6100 ft, 6; Big SpearBsh Canyon, 9 Hills. I in with am agreement Long iTii S, 3 mi W Spearfish, 5000 ft, 2; Little 2 10 mi .5800 (loc. cit.), who assigned masked shrews Spearfish Canyon, mi S, W Lead, ft, 2; 3 mi W Nemo, 4800 ft, 2; 2 mi W Nemo, from northeastern Wyoming to S. c. hay- 4700 ft, 20; Nemo, 4700 ft, 1. Pc7inington deni and those from elsewhere in that County: 3 mi W Rapid City, 1; 20 mi N Elk state to S. c. cineretis. In most dimen- Mountain, 1 (USNM); Moon, 22 mi W Hill

Table 5.—Geographic \ariation in selected color, and external and cranial measurements of adult Sorex cinereus from the Northern Great Plains.

i-i Number and sex o f«^ of c u ^ be specimens o c averaged H o TURNER: MAMMALS OF THE BLACK HILLS 43

City, 6200 ft, 1; Ditch Creek, 14 mi W Hill caverns and lesser caves have formed City, 6400 ft, 1; Palmer Culch, 3 mi SE Hill naturally in the soluble limestone layers. Citv, 53-5400 ft, 16 (U\LMZ); 3 mi SSE Hill Jewel Cave, Wind Cave, City, 1 (UMMZ); Willow Creek, 4 mi SE Davenport Ice Bear Hill Citv, 53-5400 ft, 12 (UMMZ); Castle Cave, Cave, Trap Cave, Igloo

Creek, 6500 ft, 3 ( UNLMZ); Sprins? Creek, 2 mi Cave, and the numerous unnamed and/ 5500 3 W OreviJle, ft, (UMMZ); Rapid Creek, or commercial caves of the region serve 1.5 mi W Rochford, 1 Bea\er (UMMZ); Creek, as hibernacula in the colder months. 4 mi N, 10.5 mi \V Deerfield, 6400 ft, 6; 3 mi in the Hills with the 1 N, 7 mi W Deerfield, 6400 ft, ; 3 mi S, 1 mi Mining began gold W Rockerville, 1; 16 mi NW Custer, 1 rusli of tlie mid-l

shot 8:30 P.M. be common locally (Turner and Davis, male was about (MDT) 1970:360), although previously this spe- on 9 June 1965 over Porcupine Draw; cies had been considered to be rare. this individual was found with two M. Ei^ht males were taken in a mist net set hicifugiis, one M. leihii, 11 M. volans, within the entrance to Jewel Cave on 24 and one Lasiunis cinereus. Procupine enters Ditch Creek from July (see account of M. lucifiigus). Draw Valley hillsides are clad Eight males and one female were netted the east; its steep rocky at the entrance to Ice Cave on 26 July in ponderosa pine and white spruce. five (see account of M. thysanodes), and A single, lactating female was ob- males and one female were captured in tained in Ice Cave on 26 July; two other nets set over two small ponds near Moon lactating females were taken near Moon on 30 July. An additional female was on 30 July and 20 August. A subadult netted over one of these ponds on 20 male (evidenced by unfused phalangeal Ex- August (see account of L. horealis). epiphyses) was captured at Ice Cave on rather than tensive use of mist nets, 26 July. The testes of a male shot in contrib- shooting at dusk, undoubtedly early June were 2 in length, whereas uted to increased success in capturing those of 21 males collected in late July this bat. At one place or another, all nine had an average length of 4.8 (3-6). other species of bats in the Hills have Molt was underway on 10 males taken in with Keen's been netted company in late July, but not on 11 other speci- myOtis. mens taken in the same period. A fe- The status of M. keenii in the Black male obtained on 20 August was in Hills in winter months remains uncertain. complete new pelage. The ears of the were in- Specimens have been taken in the area male shot in Porcupine Draw from 9 June through 25 August. R. A. fested by chiggers, Leptotromhidium

more . Martin (pers. com.) investigated myotis ( Ewing ) than 100 caves and mine shafts in Law- The taxonomic status of Keen's my- rence, Pennington, and Custer counties otis in the Black Hills merits further in- in winter of but M. keenii the 1969-70, vestigation. I concur with the opinion was not observed in of these hiber- 1967b: 185-186) any of Jones and Genoways ( nacula. Presumably the species repre- that individuals from the Hills fall within the Black sents a disjunct population in the currently understood range of vari- of Hills region; the nearest localities ation of M. k. septentrionalis from Kan- at Fort record are 252 miles to the north sas, Nebraska, Iowa, and eastern South Buford, Williams Co., North Dakota Dakota (Tables 6 and 7). Although and (Miller and Allen, 1928:106-107), there are no significant sexual differences 296 miles to the southeast at a locality 3 among Black Hill specimens, females mi Bon all ex- SW Springfield, Homme Co., are slightly larger than males in South Dakota (Jones and Genoways, ternal and cranial measurements. The of 1967b: 185). Surely, if this species pelage of females is also slightly paler in bat were a transient, it would be in evi- tone of color and more intense in re- dence along routes of migration. Thus, flectance of reds and blues; whereas the the based on distributional data, species pelage of males reflects greens more in- is to resident in the Hills assumed be tensely. The average breadth of the throughout the year. zygomatic arch, braincase, upper molars, Ecological information pertinent to and mastoid (excluding specimens from M. keenii in the Black Hills is limited. Iowa and Kansas) is narrower in indi- Known altitudinal range is between 4000 \'iduals from the Hills than in specimens and 6500 feet. Three specimens collected from areas to the east and southeast. on 19 July 1951 near Buckhorn, Weston Moreover the forearms of the specimens Co., ^^^'()ming, were found roosting in from the Hills are slightly shorter (ex- old buildings of a sawmill. A foraging cluding those from eastern South Da- TURNER: MAMMALS OF THE BLACK HILLS 45

Table 6.—Geographic variation in selected external and cranial measurements of adult Myotis keenii from the central United States.

Number and se.x of specimens averaged 46 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Table 7.—Geographic variation in coloration of the mid-dorsal pelage of adult Mijotis keenii obtained in .summer from the central United States.

Number and sex of specimens averaged TURNER: MAMMALS OF THE BLACK HILLS 47

Cambria Coal Mine were entering the July 1945 carried an embryo that was 13. shaft when netted between 21:15 and Testicular lengths of 17 males obtained 22:25 hrs (MDT). While collecting in from mid-June to late July were 3 or 4, Little Spearfish Canyon in late July and whereas those of three November-taken early August 1967, I located a well-used males were 2. The two males from Moon night roost abox'c Tinion Campground. and many of the specimens from Jewel One entrance to the cave was about three Cave were subadults in conspicuous molt. feet in diameter, but a second entrance, Small-footed myotis from the Black 10 by 15 feet, opened out over the sheer Hills are parasitized by a chigger, Lepto- 200-foot cliff. Most of the cave was open, twmbiclium myotis (Ewing), and a fe- roomy, and well lighted; however, to male from Jewel Cave harbored a bat tlie rear, an elongate, narrow chimney bug, Cimex piloseUus (Horvath). rose about 40 feet above the ceiling. Specimens examined (66).—SOUTH DA- Although the cave often was visited dur- KOTA: Lawrence County: Cleopatra Mine, 1.5 mi 2 Little ing the day, only one bat (a male P. N Carbonate, (SDMT); Spear- fish Canyon, Savoy, 8 nii W Lead, 5200 ft, 1; townsendii) was taken; nevertheless, at Little Spearfish Canyon, 2 nii S, 10 mi W night several bats were seen clinging to Lead, 6000 ft, 3; 2 mi W Nemo, 4700 ft, 1. walls of while the the chimney many Meade Cuuntij: Davenport Cave, 3 mi S, 0.5 others flew in and out of the large en- mi W Sturgis,' 4400 ft, 5 (NRW). Pennington 8 mi ENE 1 trance overlooking the cliff. One male County: Rapid City, (UMMZ); Rapid City, 1 (SDMT); Dark Canyon, 2 mi M. leihii, five M. hicifugus, and 10 M, S, 5 mi W Rapid City, 3800 ft, 1; Diamond S volons were collected in the roost at this Ranch, near Rapid City, 1 (UMMZ); Moon, time (Turner and Jones, 1968:444). On 22 mi W Hill City, 6200 ft, 2; Porcupine Draw, 14 mi Hill 6400 3 30 July 1968, Wayne H. Davis and I W City, ft, 1; Igloo Cave, mi S, 16 mi W Hill City, 2 (SDMT); T. 2 returned to the roost. We spent the night S, R. 6 E, E ]2 sec. 19, 1 (MSWB). Custer within the cave and established a net County: 5.75 mi N, 5.75 mi E Custer, 5220 across the two additional angling cavern; ft, 1; Roby Springs, 4 mi N, 22 mi W Custer, male M. leihii were taken, along with 24 5400 ft, 1; Hell's Canyon, 13 mi W Custer, 3 1 French Creek M. hicifugus and 42 M. volans (Turner (USNM); Custer, (AMNH); Cave, Custer State Park, 4 (SDMT); Jewel and Davis, 1970:361). Cave, 2.5 mi S, 12 mi W Custer, 5280 ft, 20 Although widespread throughout the (1 NRW, 1 SDMT, 4 UK, 1 UW); S and G Hills, populations of the small-footed Cave, 11 mi SW Custer, 1 (SDMT). Fall River 0.5 mi 1.5 mi W myotis seemingly are relatively small. County: S, Minnekahta, 4200 ft, 5. However, on 24 July 1968, Davis, Roger WYOMING: Weston County: Cambria and I 48 individuals Barbour, captured Coal Mine, 6 mi N Newcastle, 6000 ft, 8. (five females and 43 males) in a net set Additional record.—SOUTH DAKOTA: back within the entrance of Jewel Cave; Custer County: Wind Cave National Park 35 of these were banded and released (Peck, 1964:38). account of M. (see hicifugus). Thus, Myotis lueifugus carissima Thomas small-footed at myotis may congregate Little Brown Myotis favorable sites in the Black Hills. Myotis (Leuconoe) carissima Thomas, 1904, Females of this saxicolous species Ann. Mag. Nat. Hist., ser. 7, 13:383, May bear a usually only single young. Jones ( type locality, Yellowstone National Park, and Genoways (1967b: 187) reported Wyoming ) . that no embryos were found in the South Myotis lueifugus carissima—Gary, 1917, N. Amer. 3 October. Dakotan specimens examined by them Fauna, 42:43, and that most individuals collected in Myotis hicifugus is distributed the summer were males. Of three fe- throughout the northern half of North males obtained near Minnekahta in mid- America and evidently widespread and June 1967, one contained an embryo that fairly common in the Black Hills. The was 8 in crown-rump length. A female known altitudinal distribution of this taken northeast of Rapid City on 17 species extends up to 6900 feet. With 48 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

the exception of three individuals ob- bluegrass meadow 1 mi E Nemo along tained in mid-x\ugust and one taken in with one Lasiuius cinereus, one Lasion- early September, all specimens examined ycteris noctivigans, and one Eptesicus were collected in June and July. Only fuscus; one was netted on the same date two females (from Crook County, Wy- over Boxelder Creek, in Lawrence oming) taken on 1 July 1947 and 24 County along with seven L. cinereus and August, 1913, were among the specimens one E. fuscus; one was shot as it flew studied, although Jewel Cave has been over a reservoir at Roby Springs on 6 reported as a hibernaculum for both September 1968; two more were netted sexes (Jones and Genoways, 1967b: 187). over a \\'oodland pond at Moon, one each on 30 20 1968 see In the summer, females ordinarily July and August ( segregate in nursery colonies; this may account of L. borealis); another was account for their scarcity at this time. netted on 26 July 1968 at the entrance On 4 July 1970, Robert A. Martin (pers. of Ice Cave (see account of M. thy- com.) set a mist net between two chim- sa nodes). neys in the attic of the Bob Marshall On 13 June 1965, members of a field Camp dining hall, located on the shores party from the Museum of Natural His- of Bismark Lake, about 1 mi E Custer. tory obtained 12 M. lucifugus, five M. In the morning, the net yielded seven volans and a mummified Plecotus town- adult females, one subadult male, and a semlii from Jewel Cave. The majority newly born young. Martin estimated of the Myotis were located in the that 100 to 150 adults comprised the "dungeon" portion that was wet with nursery colony. To my knowledge, this seepage water and had an ambient tem- is the first report of a maternity colony perature of 6.7°C; all were torpid. Five of M. lucifugus in the Black Hills region. M. lucifugus, 48 M. leibii, eight M. Martin also captured two females in keenii, 10 M. volans, one E. fuscus, and Rapid City in the summer of 1970. four P. townsenclii were netted at the Mijotis lucifugus carissima contrasts entrance to Jewel Cave on 24 July 1968; with M. I. lucifugus of eastern South most of the bats were entering the cav- Dakota in being conspicuously more pale ern at the time of capture, between dusk and in having larger average external and and midnight. Fi\'e little brown myotis cranial dimensions (Jones, 1964 :(S6 and were caught at a night roost in Little Table 5). The subspecific name carissima Spearfish Canyon in late July of 1967 was assigned to specimens from the Wy- (see account of M. leibii), and another oming portion of the study area by Miller 24 (of which 14 were banded and re- and Allen (1928:52), and later to speci- leased) were netted there on 30 July mens from South Dakota by Jones and 1968. Genoways (1967b: 187). The average testicular length of 25 Many little brown myotis collected males taken in June and July was 4.2 in the course of this study were collected (2-7). A female, with a young attached, (shot or netted) while the bats were was extracted from a crack in the wall of foraging over water, grassy meadows or a summer cottage 15 mi ENE Sundance, along the edge of stands of ponderosa Wyoming, on 1 July 1947. pine or white spruce; several individuals Molt was apparent in an adult male were netted at cave entrances as well. taken on 7 July 1965, in Pennington Some typical situations in which M. County, because new pelage was ob- lucifugus were obtained are as follow: served underlying the old in the mid- in early July 1965 seven individuals were dorsal region and behind the ears. Sev- collected, along with two M. volans, over eral individuals from Little Spearfish a beaver pond near Beaver Creek Camp- Canyon, obtained on 30 July, were sub- ground, where the greatest activity oc- adults in conspicuous molt. An albinis- curred just after dark: two were shot tic little brown bat was captured in Jewel on 3 Jul)' 1967 over a stock pond in a Cave on 31 August 1966 by L. N. Brown TURNER: MAMMALS OF THE BLACK HILLS 49

and associates of tlie University of Wy- a subspecies restricted to the Black Hills oming. and adjacent areas. The range of this A male taken at Moon on 20 August subspecies is disjunct from the more western of M. t. 1968 was parasitized externally by bat range thysanodes by 400 miles and flies, Bosilia forcipata Ferris, and by lae- approximatelv (Jones 1967b: In laptid mites, Macronyssus crosbtji Genoways, i967a:234; 188). South has (Ewing and Storer). Mijotis lucifugus Dakota, Myotis thysanodes caiissima from many localities in the been taken only from Pennington, and Fall River Hills harbored chiggers, Leptotrom- Custer, counties, and in from Weston Its bidiinn myotis (Ewing). In a report filed Wyoming only County. at Wind Cave National Park, Olaf E. known altitudinal range in the Hills is between 3800 and R\'berg recorded another species of bat appro.ximately 6200 feet. fl)', Trichobiiis corynorJiiniis Cockerell, on specimens taken at Jewel Cave in Thirty years intervened between the May 1962. first report (Bole, 1935:147-148) and the second Spc'citncns examined (84).—SOUTH DA- report (Thompson, 1965:37) KOTA: Lawrence County: Little Spearfish of this species in the Hills region. In the Canyon, Savoy, 8 mi W Lead, 5200 ft, 3; Little interim, a young female from Custer Spearfish Canyon, 2 mi S, 10 mi W Lead, 6000 State Park (Stebler, 1939:389) and an ft, 15; 2 mi W Nemo, 4700 ft, 4; 3 mi E Nemo, adult male from 1.5 mi E Buckhorn, 4700 ft, 1; 1 mi E Nemo, 4700 ft, 3. Meade Wy- Coiiniy: Haven Dams, 6 mi S, 1.5 mi W Stui- oming (Long, 1965:532) were reported gis, 4600 ft, 1. Pennington County: Rapid as Myotis evotis. City, 1 (NRW); Sitting Bull Cave, 1 (SDMT); The fringe-tailed myotis resides in Beaver Creek, 4 mi N, 10.5 mi W Deerfield, the Black Hills the year around, as evi- 6400 ft, 8; 3 mi N, 7 mi W Deerfield, 6900 denced a male taken in ft, 1; Moon, 22 mi W Hill City, 6200 ft, 2 by hibernating (1 UK); Ditch Creek, 14 mi W Hill City, a cave along Boxelder Creek near Dody 6400 T. 2 R. 6 E M sec. 1 ft, 2; S, E, 19, Spring, in Pennington County, on 17 (MSWB); 10.5 mi E Wyoming border, 5.8 mi February 1968 (Turner and Jones, 1968: N Pennington-Custer county line, 6 (MSWB). Methods of of Custer County: 5.75 mi N, 5.75 mi E Custer, 445). capture specimens collected are Indi- 5220 ft, 1; Roby Springs, 4 mi N, 22 mi W previously unknown. 5400 2.5 Custer, ft, 1; Jewel Cave, mi S, 12 viduals obtained in the present study mi W 5280 ft, 27 8 Custer, (2 SDMT, UW, either were shot or netted as they for- 5 UK); Housing Area, Wind Cave National aged in the following areas: 1) over Park, 4100 ft, 2 (WCNP). Fall River County: water—one female over a 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 1. (5 July 1966) WYOMING: Crook County: Sand Creek, stream 3 mi N Hot Springs; one female 3750 2 15 mi ft, (USNM); ENE Sundance, (13 July 1967) over a stock pond in a 3825 ft, 1. pine-filled draw near Minnekahta; one . — Additional records ( see te.xt also ) SOUTH male (30 and two females DAKOTA: Lawrence County: Spearfish (BB). July 1968) (30 July and 20 August 1968, respec- Myotis thysanodes pahasapensis tively) over a woodland pond near Moon Jones and Genoways (see account of L. borealis; 2) over a Fringe-tailed Myotis grassy meadow—a male (19 July 1951) at dusk 1.5 mi E Buckhorn in Myotis thysanodes pahasapensis Jones and feeding Genoways, 1967, Jour. Mamm., 48:231-235, company with one Eptesicus fiisciis; 3)

20 May ( type locality, 6 mi N Newcastle, at the entrances of caves—one female 6000 ft, Weston Co., Wyoming). and five males (July and August, 1965- On 2 July 1965, I removed a fringe- 66) and one male (17 February 1970) tailed myotis from a net stretched across at Jewel Cave (see account of M. hici- a horizontal shaft in the deserted Cam- fugtis); two males (26 July 1968) at Ice bria Coal Mine complex, 6 mi N New- Cave (see account of E. ftiscus); 4) at castle, Weston Co., Wyoming. This male the opening of a mine shaft—the holo- subsequently was designated as the holo- type of pahasapensis (taken along with type of Myotis thysanodes pahasapensis, M. lei])ii). In addition, three feniah^s 50 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY were taken in mid-June 1968 while in a crest, a slightly higher and more inflated day roost, which was situated among braincase, and somewhat more robust tlie rafters of an open porch in Wind teeth (see Jones and Genoways, 1967a: Cave National Park; these fringe-tailed 233-234; 1967b: 189). Although M. evo- myotis were clustered in close association tis occurs over much of Wyoming and with M. volans (Turner and Davis, 1970: western South Dakota, no specimens 361). have been obtained in the Black Hills. When Wayne H. Davis and I netted Jones and Genoways (loc. cit.) sug- the entrances to Ice Cave on 26 July gested that thysanodes and evotis might 1968, a flurry of activity occurred be- be ecologically separated. hrs and from the Black tween 21 and 22 ( MDT), gradu- Fringe-tailed myotis ally decreased thereafter. Nine M. keenii, Hills differ from M. t. thysanodes in one M. lucifugus, 20 M. volans, and four having larger ears, shorter forearms, E. fuscus were captured with M. thy- more contrast in color between dorsal sanodes. The large mass of natural ice pelage and membranes, and a smaller in sum- skull that is narrow that persists in the cavern, even relatively ( Jones and mer, is indicative of the cool tempera- Genoways, 1967a: 233). After prelimi- tures therein. nary analysis for secondary sexual vari- Each of the three females obtained ation, the pelage of individuals collected at Wind Cave National Park on 15 June in the summer from east-central Cali- carried a single embryo; they were fornia (Tuolumne County) and from 6, 8, and 12 in crown-rump length. Lac- Cocomino, Cochise, and Gila counties, tating females were taken at Moon on Arizona, was compared with that of rep- 30 July and 20 August. A male also cap- resentatives from the Black Hills. Cali- tured on 30 July had testes that were 5 fornian specimens are significantly darker in length. Young of the year were noted in color tone than those from either Ari- flying as early as 10 August. zona or the Black Hills. Major differ- New hairs were found beneath the ences between the latter two populations old pelage of males taken in late July. are the greater reflection of greens by Molt had not begun on a female taken representatives from the Hills and the at the same time, but it was relatively slightly lighter tone of the individuals far advanced on a female obtained on from Arizona. Average measurements 20 August. Three pregnant females cap- and standard deviations of pelage color six tured at Wind Cave National Park on 15 of 10 ( four males and females ) adult June were molting over both dorsum and M. t. pahasapensis collected in the sum- venter. mer are: color tone, 25.3 ±4.22; percent A female fringe-tailed myotis from red reflectance, 55.0 ±0.56; percent green Fall River County harbored a bat fly, reflectance, 20.1 ±0.29; and percent blue Basilia forcipata Ferris, chiggers, Lepto- reflectance, 24.9 ±0.36. Selected average trombicUiim myotis (Ewing), and a bat measurements and standard deviations of bug, Cimex adjunctus Barber. Dr. George 16 specimens, nine males and seven fe- W. Byers, Department of Entomology, males, of fringe-tailed myotis from the The University of Kansas, identified the Black Hills are: total length, 94.4 ±3.22; remains of a moth, a small scarabid tail length, 41.0±1.94; hind foot length, beetle, and a large alleculid beetle from 10.6±0'!^92; ear length, 18.5±0.95; fore- the stomach of this same individual. arm length, 41. 1± 1.20; greatest length In contrast with Myotis evotis, which of skull, 16.6 ±0.27; zygomatic breadth, the fringe-tailed bat resembles superfi- 10.1 ±0.21; interorbital constriction, 3.9 cially, Myotis thysanodes possesses a ±0.14; braincase breadth, 7.7 ±0.19; longer forearm, shorter ears, slightly mastoid breadth, 8.3 ±0.19; breadth larger cranial dimensions, a conspicuous across upper molars, 6.5±0.22; maxillary fringe of hairs on the free border of the tooth-row length, 6.3 ±0.16; and breadth uropatagium, a better developed sagittal across upper canines, 3.9±0.18. TURNER: MAMMALS OF THE BLACK HILLS 51

S))ccimens examined (30).—SOUTH DA- flew through Ditch Creek Valley and KOTA: Boxclder Pennington County: along associated Porcupine Draw in Penning- Creek, near Dodv Spring, 4060 ft, 1; Moon, ton County (see account of M. keenii). 22 mi W Hill Cit\, 6200 ft, 3; Ice Cave, 8 mi Five males and two females were taken N, 15 mi W Custer, 6400 ft, 2 (UK); 10.5 mi E \\'\oming border, 5.8 mi N Pennington- in company with one M. leihii over a Custer Custer count)- line, 7 (MSWB). County: reservoir at Roby Springs, Custer Grace Coolidge Creek, Custer State Park, 1 County, in early September 1968. Five (UMMZ); jewel Cave, 2.5 mi S, 12 mi W males and two females (four of these Custer, 5280 ft, 9 (3 SDMT, 1 UMMZ, 4 UW); Ele\'ator Building, Wind Cave National Park, bats were banded and released) were 3 Hot 1 Fall 4100 ft, 3; mi N Springs, (UW). netted over ponds near Moon in late River 0.5 mi 1.5 mi W County: S, Minnekahta, July and mid-August of 1968 (see ac- 4200 ft, 1. count of L. horealis). A female, evi- \\TOMI\G: Weston County: 1.5 mi E disturbed from her roost, Buckliorn, 6150 ft, 1; Cambria Coal Mine, 6 dently daytime in mi N Newcastle, 6000 ft, 1. was captured mid-morning (10:30 A.M., MDT) at the Beaver Creek Camp- Myotis volans interior Miller ground, Pennington County, on 13 June Long-legged Myotis 1965. A male collected at this same site on 2 1965 carried the remains of a Myotis longicrus interior Miller, 1914, Proc. July Biol. Soc. Washington, 27:211, 31 October crane fly (Tipulidae) in his mouth. (type locality, 5 mi S Twining, 11,300 ft, Caves are utilized in summer by M. Taos New Co., Me.xico). volans both as daytime retreats and as volans interior—Miller and G. M. Myotis Allen, nighttime roosts following foraging Bull. U. S. Nat. 25 1928, Mus., 144:142, Individuals have been obtained May. flights. during the summer from Bear Trap Cave, Myotis volans, first reported in the Ice Cave ( 19 banded and released on 24 area is study by Moulthrop (1936:413), July 1968), and Jewel Cave. (Five tor- known the Black ex- throughout Hills, pid bats were obtained in the latter cav- from Crook Evi- cept County, Wyoming. ern on 13 June 1965; see account of M. dently this is the most common and liicifugus. Nine M. volans were banded distributed of the widely species genus and released in Jewel Cave on 24 July in the In this region. general, species 1968.) On 5 July 1967, I shot a male montane areas of western North occupies about 10:30 P.M. (MDT) as it flew into America. The known altitudinal range Davenport Cave and alighted on a crys- in the Black Hills is 4000 to 6500 feet, talline calcite (dog-tooth spar) wall. Ten but the is most abundant above species males were collected from a night roost 4500 feet. It is resident the throughout in Little Spearfish Canyon between 26 year as evidenced indi- by hibernating July and 7 August 1967 (see account of viduals taken from Bush's Cave (20 No- M. leihii), and 42 males were netted, and Cave to 26 No- vember) Jewel (20 banded, and released in this same cavern vember and 17 February). on 30 July 1968. All chiropteran species The long-legged myotis superficially known to occur in the Black Hills were resembles the little brown myotis, but associated with M. volans at some local- M. volans differs from M. in hiciftigus ity. On two different occasions, individ- having a shorter rostrum, a braincase uals were located in a day roost on Wind that is abruptly elevated from the rostral Cave National Park in company with M. level, low rounded ears, a keeled calcar, thysanodes. Eleven of these were banded and pelage that usually extends onto the and released, and a female was netted underside of the wing to a line joining one month later within 200 yards of the the knee. elbow and original site of capture (Turner and Myotis volans forages over such areas Davis, 1970:362). as campgrounds, meadows, and water On 20 November 1967, a single male courses. For example, in early June 1965, was taken in Bush's Cave, a small cave 11 individuals were shot at dusk as they several hundred feet south of the en- 52 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

trance to Jewel Cave. The ambient tem- Jewel Cave, 2.5 mi S, 12 mi W Custer, 5280 ft, 12 (3 NRW, 1 SDMT, 1 UW); Headquar- perature was 9.7 °C and the rectal tem- ters, Wind Cave National Park, 4100 ft, 1 of the bat, which was perature torpid (UMMZ); Elevator Building, Wind Cave Na- and covered of by droplets moisture, tional Park, 4100 ft, 1; Housing Area, Wind was 10.6° C. A single Plecotns townsendii Cave National Park, 4100 ft, 5. Fall River also was noted in Bush's Cave at that County: 5.5 mi E Minnekahta, 4000 ft, 1; 0.5 mi S, 1.5 mi W 4200 2. time. Minnekahta, ft, WYOMING: Weston County: 1.5 mi E

and 1967b : 190 Jones Genoways ( ) Buckliorn, 6150 ft, 1; Cambria Coal Mine, 6 reported two females, each containing a mi N Newcastle, 6000 ft, 1. Additional record.—SOUTH DAKOTA: single embryo, that were obtained on Custer County: Wind Cave, Wind Cave Na- 29 and 30 June in Harding County, north tional Park (Peck, 1964:39). of the Hills. However, of 18 females collected in the summer and examined Lasionycteris noctivagans (Le Conte) from the Hills none was lactat- region, Skiver-HAIRED Bat ing and only one was pregnant. The Le in latter specimen was taken in Wind Cave V[espertilio]. noctivagans Conte, 1831, McMurtrie, The animal kingdom ... by National Park on 25 July and the single the Baron Cuvier . . ., 1:431 (type local- was 16 in A embryo crown-rump length. ity, eastern United States). of the was ob- male flying young year [Lasionycteris] noctivagans—Peters, 1866, Mo- tained at Roby Springs on 7 September. natsb. k. preuss. Acad. Wiss., Berlin, p. 648 Testes of 14 males taken from mid-June (for 1865). to late July were 1 to 5 in length. The silver-haired bat occurs through- New hairs were manifest under the out the United States, but is known in old pelage over much of the body of a the Black Hills from only seven localities in male taken on 30 July 1961. Bat flies, South Dakota; all but two specimens Basilia forcipata Ferris, were found on were obtained in transitional areas be- M. volans from Pennington, Custer and tween the high plains and foothills along Fall River counties, whereas other indi- the southern and eastern periphery of viduals from Pennington, Fall River, and the Hills. This species first was reported Lawrence counties harbored chiggers, in the study area by Jones and Genoways Leptotrombidium mijotis (Ewing). Two (1967b: 190). Although little is known females from Wind Cave National Park of the actual migratory routes, Lasion- were infested by spinturnicid mites, ycteris presumably migrates through the Spinturnix americanus (Banks), and a Hills region (northward in spring and male from Roby Springs, Custer County, southward in late summer and early was parasitized externally by several un- autumn). Jones and Cenoways (loc. cit. that this identifiable (badly damaged) mites. ) suggested species probably is resident in the of the Black Specimens examined (77).—SOUTH DA- higher part KOTA: Lawrence County. 2 mi S Tinton, 6100 Hills in summer. The altitudes at which ft, 1; Little Spearfish Canyon, 2 mi S, 10 mi most indi\iduals ha^e been collected W 6000 10; 2 mi W 4700 ft, Lead, ft, Nemo, thus far generally range from 3800 to 1. Meade County: Davenport Cave, 3 mi S, 4700 feet, but two males were obtained 0.5 mi W Sturgis, 4400 ft, 1. Pennington at 6200 feet near Moon on 29 1968 County: Rapid City, 3 (NRW); Beaver Creek, July, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 3; Moon, (see account of L. horealis). Thus, the 22 mi Hill 6200 14 W City, ft, 3; Ditch Creek, silver-haired bat is more widely distrib- mi W Hill City, 6400 ft, 11; Ice Cave, 8 mi uted altitudinally than previously sup- N, 15 mi W Custer, 6400 ft, 1; Bear Trap posed. Cave, 4 mi S, 16 mi W Hill City, 6000 ft, 1; T. 2 S, R. 6 E, E M sec. 19, 1 (MSWB); 10.5 mi There are few records of silver-haired E 5.8 mi N Custer Wyoming border, County bats occurring in caves. A solitaiy male 8 1 line, (MSWB); unspecified locality, was taken from an unnamed cave in Dark (NRW). Custer County: Roby Springs, 4 mi on 19 November 1967 in a semi- N, 22 mi W Custer, 5400 ft, 7; Bull Springs, Canyon state. limestone ca\'e is lo- 2 mi N, 9 mi W Custer, 6500 ft, 1 (UMMZ); torpid The TURNER: MAMMALS OF THE BLACK HILLS 53 cated about 90 feet below the crest of County, was parasitized externally by a 200-foot cliff. Relative humidity within chiggers, Leptotrojnhidium myotis the cave was 57 percent, and the ambient (Ewing), whereas the male from Dark Mac- temperature was 6.7 ""C; rectal tempera- Canyon harbored laelaptid mites, true of the bat was 9.6^C. The male was ronyssus tinidens Radovsky. In addition, hidden in a horizontal crevice and be- B. V. Peterson, Entomology Research a came somewhat acti\'e shortly after being Institute, Ottawa, Ontario, identified extracted. Three Plecotus townsendii bat fly found on a male L. noctivaiians Park as either and one Myotis leibii, all in a torpid from Wind Cave National or a state, also were collected in the same Trichohitis adamsi Auguston new cavern at this time. The lateness of the species; T. adamsi has been reported season, the low rectal temperature, and previously only from Macrotus californi- tlie situation in which the bat was found, cus and Tadarida hrasiliensis. this was not a all suggest that individual Specimens examined (11).—SOUTH DA- late transient, but rather in hibernation KOTA: Lawrence County: 2 mi W Nemo, 4700 1 mi E Nemo, 4700 ft, 1. Penning- (Turner and Jones, 1968:445). Thus, it ft, 2; ton County: Diamond S Ranch, near Rapid is probable tliat at least a few silver- City, 1 (UMMZ); Dark Canyon, 2 mi S, 5 mi haired bats reside in the Black Hills W Rapid City, 3800 ft, 1; Moon, 22 mi W the entire throughout year. Hill City, 6200 ft, 2. Custer County: Wind In 1967, five individuals, two from Cave Canyon, Wind Cave National Park, 4100 1. Fall River 0.5 mi 1.5 mi W southwest of Minnekahta (12 and 13 ft, County: S, Minnekahta, 4200 ft, 3. June) and three from near Nemo (29 to 3 were shot as June July ) , they foraged Eptesicus fuscus pallidus Young in with in early evening company Myotis Big Brown Bat leibii, M. hicifugtis, M. thysanodes, Ep- Eptesicus pallidus Young, 1908, Proc. Acad. tesicus fuscus, and Lasiurus cinereus. Nat. Sci. Philadelphia, 60:408, 14 October Another individual was at the captured (type locality, Boulder, Boulder Co., Colo- site at 13:30 on 14 fonner (MDT) June rado ) . in a net over a small algae-covered pond, Eptesicus fuscus pallidus—Miller, 1912, Bull. and a male was captured in a net set U. S. Nat. Mus., 79:62, 31 December. over a at the head sewage settling pond The pallid big brown bat, originally of Wind Cave on 25 1968. Canyon July reported from the region as Adelonyc- This seems to to species prefer forage teris fusca (J. A. Allen, 1895a:273), is that contain a source in grassv vallevs common, widespread, and occurs in all water and which are sur- of standing counties that comprise the Black Hills. rounded hillsides well-forested with by This species inhabits most of North p'jnderosa pine. America and is known in the study area Females give birth to one or usually from elevations between 3500 and 6200 two young between late May and early feet. Eptesicus fuscus pallidus differs shot Minnekahta is July. A female near from E. f. fuscus which geographically on 12 June 1967 carried two embryos adjacent to the east in South Dakota in that were 11 in crown-rump length. its discernibly paler pelage, and smaller Length of the testes of five males taken average external and cranial dimensions from 13 June to 3 July 1967 varied be- (Jones, 1964:92). tween 4 and 5; a male obtained in No- Most of the specimens examined were vember had a testicular length of 4. The collected in summer; however, individ- testes of three males obtained in late July uals have been obtained in the winter 1968 were 6, 7, and 7 in length. Fine at Custer (23 January 1888) by Vernon silver-tipped hairs were seen underlying Bailey, in Hell's Canyon (5 January the old pelage on the dorsum of the three 1902) by Merritt Cary, and in hibemac- latter individuals. ula such as Jewel Cave (Jones and Geno- A male from Minnekahta, Fall River ways, 1967b: 192). On 7 February 1970, 54 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Robert A. Martin and associates (pers. rump length) in the attic of a classroom com.) located a winter colony of 20 E. building on the campus of the Black fusctis, four Myotis leibii, and two Pleco- Hills State College at Spearfish. Females tus townsenclii in a cave along French ordinarily give birth to one or two young Creek in Custer State Park; all bats between May and mid-June in western found in the cavern at that time were South Dakota (e.g., a female taken in males. Fall River County on 14 June carried While attempting to obtain flying two embryos that each measured 7 in in a on 11 thus a squirrels spruce-filled canyon crown-i-ump length ) ; pregnant big February 1946, J. A. King captured a brown bat in late July would seem ex- female big brown bat by the wing in a ceptional. Returning to the same attic rat trap; the trap was attached about on 29 July I obtained 21 additional E. seven feet above the ground to a tree, fusciis, of which eight were lactating and peanuts were used as bait. The local adult females, five were subadult fe- conditions under which this unseasonal males, seven were subadult males, and activity occurred are unknown to me. one was an adult male. later collected King a male (4 March Martin (pers. com.) reported that a 1946) and a female (21 March 1946) maternity colony of this species was lo- that were hibernating in an old hori- cated on 19 May 1970 in the attic of zontal mine shaft in the same area 3 mi ( the Dakota Steel Company in Rapid City. SE Hill City, Pennington County). Two other maternity colonies were Many of the big brown bats collected found in Hot Springs in the summer of in this study were shot at dusk as they 1968. Ten pregnant females (six were foraged over grassy, conifer-bordered banded and released, and four were ex- meadows, or over ponds and water amined; each of the latter carried a courses (Turner and Jones, 1968:446). single embryo of an average crown-rump Others were captured in mist nets in the length of 18.5 (15.0-23.0)) and four lac- following situations: One male (3 July tating females were netted in the attic 1967) captured as it flew along Boxelder of the Fall River Museum on 25 June Creek, 2 mi W Nemo (see account of and 27 July, respectively. One subadult

M. lucifu

Four males and one . less Myotis lucifiigus). 363 ) The entire process lasted than female (26 July 1968; four banded and 10 minutes. The female propped herself released) netted at the entrance of Ice on the tips of her forearms, lowered her Cave. The main activity of 32 bats taken head ventrally toward the vaginal ori- in the last-mentioned cavern was be- fice, and actively aided the birth by 21 tween and 22 hrs (MDT); however, licking the vaginal region and the emerg- big brown bats were not obtained until ing young. The young were cradled in about 23 hrs (see account of Myotis the uropatagium, which was curved for- thysanodes) . Other species taken along ward beneath the body of the female. with Eptesicus at various sites of captiuc^ Though seemingly close to exhaustion, were Myotis keenii, M. leihii, M. hici- the female readily devoured several fugus, M. thysanodes, M. volans, Lasi- moths offered to her immediately follow- onycteris noctivagans and Lasiurus cine- ing the events just described. reus. Length of testes of 18 adult males ob- On 27 July 1967, I captured a female tained between 13 June and 4 July were that carried two embryos (8 in crown- 5.8 (4.0-7.5). Testicular length of two TURNER: MAMMALS OF THE BLACK HILLS 55

1 Weston 1.5 mi E Buck- adults taken on 29 Jul)' 1967 were 9 and (USNM). County: horn, 6150 ft, 1; Cambria Coal Mine, 6 mi N 10, whereas that of five subadults cap- Newcastle, 6000 ft, 1. tured on the same were 4.4 day (3.0-7.0). Additional record.—SOUTH DAKOTA: A male (banded and released), with Meade County: Fort Meade (Miller, 1897:98). large, descended testes, was extracted from a space between a stone chimney Lasiurus borealis borealis (Miiller) and eaves of an overhanging roof of a Red Bat house in Rapid City on 3 August 1969. Vespertiliu borealis Miiller, 1776, Des Ritters Jones and Genoways (1967b: 191) Carl von Linne . . . vollstandiges Natm- described molt in E. and their f. juscus system . . ., suppl., p. 20 (type locality.

to E. York . description seems applicable also f. New ) pallidiis. Subadults from Spearfish and Lasiurus borealis—Miller, 1897, N. Amer, 16 October. Hot Springs were undergoing post- Fauna, 13:105, juvenal molt late in July, whereas an Turner and Davis (1970:363) re- adult female from Spearfish was in proc- cently reported the first specimens of the ess of annual molt. red bat to be taken west of the Missouri Big brown bats from southwest of River in South Dakota. Jones and Geno- Minnekahta harbored bat bugs, Cimex ways (1967b: 193) suggested that this adjunctus Barber, and ticks, Ornitho- species probably reached the western doros kelleyi Cooley and Kohls, whereas limits of its resident range in South Da- a female from Hot Springs was infested kota at approximately the 101st meridian,

' with spinturnicid mites, Spinturnix hakeri but that migrating individuals might pass Rudnick, and a male from Moon shel- through the western part of the state on tered bat flies, Basila forcipata Ferris. their way to or from regions farther to Eptesicus from various localities in the the north. Long (1965:534) mentioned Black Hills were parasitized externally an individual (USNM 5264) that was by chiggers, Leptotrornbidhitn imjotis hsted by H. Allen (1864:20, 1894:153) (Ewing). from Laramie Peak, as the only specimen known from Specimens examined (129).—SOUTH DA- Wyoming. KOTA: Lawrence County: Spearfish, 3600 ft, Nets were established over two ponds Little 8 mi 22; Spearfish Canyon, Savoy, W near Moon Campground in Pennington Lead, 5200 ft, 2; 2 mi W Nemo, 4700 ft, 6; County on 29 July 1968. One net was 1 mi E Nemo, 4700 ft, 1. Pennington County: located in a small meadow in a woodland Rapid City, 5 (4 NRW, 1 SDMT); Diamond S Ranch, near Rapid City, 6 (UMMZ); South and the other over a farm pond situated 3 mi 12 Canyon, W Rapid City, (AMNH); in a pasture. No red bats were taken Moon, 22 mi W Hill 6200 ft, 1; Ice Cave, City, over the woodland pond, but Wayne H. 8 mi N, 15 mi W Custer, 6400 ft, 1; Hill City, Davis a female of the 1 (NRW); 3 mi SE Hill City, 5300-5400 ft, captured young 3 (UMMZ); 10.5 mi E Wyoming border, 5.8 year over the farm pond; the latter indi- mi N Custer County line, 2 (MSWB). Custer vidual was just beginning post-juvenal County: 5.75 mi N, 5.75 mi E Custer, 5220 molt. Two Mijotis leibii, two M. keenii, ft, 5; Hell's Canyon, 13 mi W Custer, 2 two M. volons, two Lasiurus cinereus, (USNM); Custer, 4 (USNM); Squaw [Grace and two were Coolidge] Creek, 1 (AMNH); French Creek Lasionycteris noctivagans Cave, Custer State Park, 8 (SDMT); Jewel removed from this same net, whereas 2.5 mi 12 mi 5280 2 Ca^e, S, W Custer, ft, four M. keenii, one M. lucifuf^us, two M. (1 UW, 1 WCNP); Headquarters, Wind Cave fJiysanodes, three M. volans, four E. fus- National Park, 4100 ft, 17 (2 WCNP, 15 cus and 13 L. cinereus were removed UMMZ); Shirttail Canyon, Wind Cave Na- tional Park, 2 (UMMZ); Ony.x Cave, 10 mi NW from the net over the woodland pond. Hot Springs, 1 (UW). Fall River County: Returning to the woodland pond on 20 Fall River Museum, Hot 3600 ft, 8; Springs, August, I obtained a lactating female red Courtliouse, Hot Spnngs, 3500 ft, 5; 5.5 mi E bat, along with one M. keenii, one M. Minnekahta, 4000 ft, 1; 0.5 mi S, 1.5 mi W one M. two M. Minnekahta, 4200 ft, 8. Jucifufi^us, thysonodes, WYOMING: Crook County: Sand Creek, volans and one L. cinereus. On the first 56 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

seasonal records are 13 visit, the woodland pond was fairly open autumn; from and activity began about 21 hrs (MDT); June to 31 August. a total of 27 bats was netted. In contrast, Findley and Jones (1964:469) indi- with the pond was well overgrown vege- cated that females generally precede the tation on the second visit. Although males in the northward migration and from 20 until 00:30 net was observed occupy the plains and adjacent lowlands hrs, not a single bat was detected; how- in warmer months, whereas the males the next ever, when examined early usually are found at higher altitudes or, seven bats in the net were morning, the occasionally, at higher latitudes. The 28 concentrated over a small pool of open specimens (11 males and 17 females) the bats water. Utilization of pond by taken in this study certainly establish the evidently decreased greatly as the sur- Black Hills as one of the few areas in face vegetation increased. the mid-continent region where adult L. The interim (22 days) between cap- cinereus of both sexes occur together in ture of the flying young of the year and summer. In western South Dakota, males the lactating female almost certainly ex- occur only in the Black Hills, at eleva- cludes the possibility that the young was tions of 4700 to 6200 feet, whereas fe- that of the female. Thus, at least two males occupy suitable habitat up to 6400 families of L. borealis evidently occurred feet in the Hills and also range down in the vicinity of Moon during the sum- through transitional lower elevations out mer of 1968. Members of this species onto the adjacent plains and lowlands. migrate southward in winter, returning The first representatives of this species northward again in spring, and addi- from the Black Hills were reported by

to ascertain the : 193 . tional data are needed Jones and Genoways ( 1967b ) summer status of the red bat in the Black Of 14 hoarv bats collected in the Hills. vicinity of Nemo between 28 June and 3 Specimens examined (2).—SOUTH DA- July 1967, seven were netted over Box- 22 mi KOTA: Pennin0on County: Moon, W elder Creek (see account of M. Jucifu- Hill City, 6200 ft, 2. and seven others were shot in areas gus ) , where M. leibii, M. hicifugus, M. voJans, Lasiurus cinereus cinereus E. fusctis, and Lasionycteris noctivagans de (Palisot Beauvois) also foraged (i.e., over streams and mea- Hoary Bat dows bordered by ponderosa pine). A single male was shot on 19 June 1967 as cinereus Pali- Vespertilio (misspelled lincreus) it foraged over a small valley northeast sot de Beauvois, 1796, Catalogue raisonne of Custer, and two females were obtained du museum de Mr. C. W. Peale, Phila- netted over a stock in asso- delphia, p. 18 (p. 5 of English edition liy (one pond,

Peale and Palisot de Beauvois ) ( type lo- ciation with the above mentioned species cality, Philadelphia, Pliiladelphia Co., Penn- and M. Hu/sanodes, and the other shot as sylvania). — it flew along the edge of a conifer- Lasiurus cinereus H. Allen, 1864, Smiths. Misc. bordered meadow) southwest of Minne- Coll., 7 ( 165 ):21, June. kahta on 13 June 1967. Of 15 hoary bats The bat has not 1 1 released netted over hoary been reported ( banded and ) from the Wyoming section of the Black a woodland pond (13 individuals) and Hills, but it is known from Lawrence, a farm pond (two individuals) near Pennington, Custer, and Fall River coun- Moon on 30 July 1968, six were adult ties of South Dakota. This is the largest males, six were lactating females, and of the Black Hills bats and seems to be three were juveniles (one male and two locally abundant in suitable habitats. females); an additional male young of Like the red bat, Lashinis cinereus is a the year was obtained o\'er the wood- migrant, arriving in the Hills region in land pond on 20 August 1968 (see ac- late spring and residing there until early count of L. borealis). Two female L. TURNEH: MAMMALS OF THE BLACK HILLS 57 cinereus (one lactating and the other a 2000 big-eared bats hibernate in winter young of the year) were netted over a along with Mijotis leihii, M. liieifugus, sewage setthng pond at the head of M. volans, and Eptesieus fiiscus. Bats Wind Cave Canyon on 25 July 196(S in evidently vacate these limestone hiber- company with a L. noctivag,ans. nacula in late spring and disperse One or (usually) two young are born throughout the Hills; females congregate from late May to early July. Of two in maternity colonies in warm standstone adult females taken on 13 June, one car- caves or attics of buildings, whereas ried two embryos (each being 22 in males seem to remain solitary. Known crown-rump length) and the other was altitudinal range in the Hills is from 3400 lactating. None of six females taken be- to 6200 feet. tween 28 June and 3 July was pregnant, Although large associations of big- but four were lactating; all six adult eared bats occur in the Hills region, none females obtained at Moon on 29 July, of the specimens taken in this study was and one collected in Ditch Creek Valley obtained by the usual collecting methods, on 13 also Testes June, were lactating. that is, by netting or shooting. Perhaps of eight males captured from mid-June the activity patterns of this bat differ to mid-July were 5.4 (4-7) in length. from those of other species that occur The subadults collected late in July in the region, and it may remain in re- to mid-August were in fresh pelage. No treats until well after dark. V. Bailey, ectoparasites were found on individuals however, reported P. townsendii to be of this species. among the earliest observed bats flying Specimens examined (28).—SOLTTH DA- each evening in the vicinity of Sundance, KOTA: Lawrence County: 2 mi E Tinton, Wyoming (Handley, 1959:175). 5400 ft, 1; 2 mi W Nemo, 4700 ft, L3; 1 mi On 4 August 1969, a solitary big-eared E Nemo, 4700 ft, L Pennington County: bat was observed in a narrow Rapid Citv, 1 (NRW); Moon, 22 mi W Hill chimney Cit>-, 6200 ft, 5 (2 UK); Ditch Creek, 14 mi of a small cave on the rim of Little Hill 6400 10.5 mi E W City, ft, 1; Wyoming Spearfish Canyon, above Timon Camp- border, 5.8 mi N Pennington-Custer county ground; a single male (testes 6 in length) line, 1 (MSWB). Custer Counti/: 5.75 mi N, was in the cave on 7 5.75 mi E Custer, 5220 ft, 1; Wind Cave Can- captured August 1970. the is used exten- yon, Wind Cave National Park, 4100 ft, 2. Although cavern Fall River Countti: 0.5 mi S, 1.5 mi W Minne- sively as a night roost (see account of kahta, 4200 ft, 2. M. leihii), P. t. palleseens were the only bats that utilized the chimney as a day Piece tus townsendii palleseens (Miller) roost. Townsend's Big-eared Bat On 20 July 1967, I located a nursery Corynorhinus macrotis palleseens Miller, 1897, colony in a small cave formed in a sand- N. Amer. Fauna, 13:52, 16 October (type stone bluff at a locality 1 mi S and 2 mi locality, Keam Canyon, Navajo County, Ari- zona). E Hot Springs, Fall River County Plecotus townsendii palleseens—Handley, 1959, (Turner and Jones, 1968:447). Of the Proc. U. S. Nat. Mus., 110:190, 3 Septem- 41 bats present in the cavern, 22 were ber. adult females; 19 of these were lactating This cavernicolous species is distrib- and one carried an embryo that was 27 in uted through much of western North crown-Runp length. Eight juvenile fe- America, and is known from all counties males and 11 juvenile males (with tes- that comprise the Black Hills; it was first ticular lengths of 3) comprised the re- reported from the study area by G. M. mainder of the colony. The young rep- Allen (1916:334) under the name Cory- resented various stages of development norhinus megalotis palleseens. John Ty- and molt (from fine post-natal hair on ers and associates of the U. S. National nearly naked animals to juvenal pelage Park Service have banded many individ- on well de\'eloped }'oung) indicating uals in Jewel Cave, where an estimated that dates of parturition within the col- 58 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY ony probably extend over a considerable vaginal orifice was observed in many of period of time. A solitary male (testes these females when rectal temperatures 6 in length) was found in this same cave were taken. Thomas H. Kunz examined on 30 June 1968. seven vaginal smears under a phase con- A secluded male (testicular length 4) trast microscope and found numerous was found clinging to the crystalline cal- agglutinated clumps of sperm in the cite walls of semen Pearson et (dog-tooth spar) Daven- suspension. ah ( 1952: port Cave on 20 November 1967, and 293) indicated that vaginal plugs were three bats not found in Plecotus big-eared ( two males and one townsendii studied female) were found hibernating in an in California. unnamed limestone cave in Dark Can- Jones and Genoways (1967b: 194) yon on 10 November 1967 (see account noted the complete absence of big-eared of L. noctivagans) . Although the males bats in Jewel Cave on 12 June 1965, and were captured deep in the cave, the speculated that Plecotus vacates the cave solitary female was taken in the twilight entirely in late spring and for most of the zone. Testicular length of both males was summer. However, nets placed within 3. Ambient temperatures at the sites of the entrance to Jewel Cave on 24 July capture were 6.4°, 5.3°, and 7.2 °C, 1968 captured four Plecotus along with whereas the rectal temperatures of the several other species (see account of M. bats were 7.2°, 8.0°, and 8.6°C, respec- hicifugus). Of the 2165 big-eared bats tively. Mummified remains of P. toicn- banded by the National Park Service at sendu were found in Jewel Cave on 13 Jewel Cave from December 1959 through June 1965, and in the deserted Cambria December 1963, nine returns have been Coal Mine complex north of Newcastle, reported. Four of these were taken Weston County, Wyoming, on 2 July within an area 30 miles south and east 1965. of Jewel Cave between elevations of 3400 Jewel Cave was visited by a field and 3800 feet (see additional records) party from the Museum of Natural His- during the warm months (5 May to 1

. The others were tory on 20 November 1967. Approxi- September ) recaptured mately 600 Plecotus were wintering within the cave itself in winter. A male there at the time, and 304 of these were that was banded on 31 December 1959 examined. Two hundred big-eared bats was recaptured in Jewel Cave on 24 July (69 females and 131 males) were distrib- 1968; thus, there was an interim of 3148 uted in clusters containing from two to davs between the time of banding and 33 bats; the remaining 104 individuals recapture (Turner and Davis, 1970: 40 females and 64 males ( ) were solitary. 364). Ambient temperature ranged from 5.0° Of those specimens examined in to 6.4°C and the relative humidity Jewel Cave on 20 November 1967, 67 from 64 to ranged 70 percent in areas percent were parasitized by bat flies, occupied by Plecotus. The average rec- Trichohius corynorliini Cockerell and tal of temperatures 18 females and 34 one individual harbored a chigger, Lep- males were 4.3°C and 4.2°C (3.6-6.2) totromhidium mijotis (Ewing). Sarcop- (3.6-6.2), individuals from respectively; tid mites, Sarcoptes lasionycteris Boyd larger clusters tended to have a mean and Bernstein, were prevalent also in the body temperature 1-2° lower than did Jewel Cave population. Female big- solitary individuals or those from small eared bats a incidence clusters. Most of the bats were within displayed higher of parasitism than did males. Two fe- 150 feet of the entrance, where the am- males from the near bient temperature was lower than deeper maternity colony Hot were in the cave. Air currents made this 2:en- Springs, parasitized by the eral area susceptible to environmental same species of bat fly and by laelaptid fluctuations. mites, Macronyssus longisetosus ( Fur- A that filled plug completely the man ) , whereas a female from Dark Can- TURNER: MAMMALS OF THE BLACK HILLS 59 yon harbored a different species of lae- Leptis may occur in the immediate vi- laptid mite, Macronyssus iinidens Ra- cinity of the Hills. The four native dovsky, representing tlie first record of leporids represent interesting examples this parasite in Soiitli Dakota. of similar species with either parapatric One of tlie 22 adult females from the or sympatric, but ecologically distinct, nursery colon) near Hot Springs (ob- distributions in the Hills region. tained on 20 July 1967) was molting audubonii simultaneously over both dorsum and Sylvilagus baileyi (Merriam) venter. Desert Cottontail examined SOUTH DA- Specimens (168).— Lej)us baileyi Merriam, 1897, Proc. Biol. Soc. KOTA: Lawrence Coiuitt/: Mine, Cleopatra Washington, 11:148, 9 June (type locality, 1.5 mi N 2 T. 5 R. Carbonate, (SDMT); N, Spring Creek, E side Bighorn Basin, Big 2 NE }, sec. 2 "near Chicken E, 17, (SDMT); Horn Co., Wyoming). Ranch and Rouliaix Lake," 1 (NRW); Crow- Sylvilagus auduhonii haileyi—Lantz, 1908, feet [=Cro\v Peak?], Black Hills, 7 (AMNH); Trans. Kansas Acad. Sci., 22:236. R. B. Hayes Mine, 1 (SDMT). Meade County: Da\enport Cave, 3 mi S, 0.5 mi W Sturgis, Nelson (1909:234) first reported the 4400 ft, 1. Pennington County: Tool Shed desert cottontail from the Black Hills on Cave, N Battle Creek, S Haywood, 3570 ft, 8; the basis of a young female obtained at Rochford, 5300 ft, 1 (UMMZ); Deadman Can- Elk Mountain Merritt on 10 yon, 2-3 mi W Rapid City, 1 (NRW); Rapid by Gary City, 1 (NRW); Dark Canyon, 2 mi S, 5 mi W July 1906. Sij]vi]a(i,us auduhonii is a Rapid City, 3800 ft, 3; Spring Creek Canyon, common mammal on the western Great 1 (NRW); 4 mi SW Rockerville, 5000 ft, 1 Plains and inhabits grasslands up to (UMMZ); Igloo Cave, 3 mi S, 16 mi W Hill about 5000 feet in the Hills. Cit>-, 12 (SDMT); 3 mi SE Hill City, 3 Specimens are available (UMMZ); Bear Trap Cave, 4 mi S, 16 mi W only from Pennington, Cus- Hill 6200 2 1 Custer City, ft, ( SDMT). County: ter, and Fall River counties, although Hell's 13 mi W Custer, 7 (USNM); Canyon, the species undoubtedly occurs along the Mine, 2 (SDMT); 18 mi W Custer, Cuvahoga entire interface of the Black Hills and 1 (USNM); Jewel Cave, 2.5 mi S, 12 mi W The desert cotton- Custer, 5280 ft, 52 (2 NRW, 3 SDMT, 1 UW, surrounding prairie. 4 WCNP); S and G Cave, T. 5 S, R. 3 E, tail is geographically sympatric with the NW }i, NW }i sec. 3, 2 (SDMT); 2 mi W Wind eastern cottontail and with Nuttall's cot- Cave National Park, 1 (UW); Wind Cave, tontail; however, these three rabbits usu- Wind Cave National Park, 2 (1 SDMT, 1 different niches. UW). Fall River County: 1 mi S, 2 mi E Hot ally occupy ecological Springs, 3400 ft, 41; 5.5 mi E Minnekahta, In the western section of the Hills, 4000 ft, 1. Nuttall's cottontail has been collected WTOMINC: Crook County: Sand Creek, in the same general area with the desert 10 mi E Sundance, 3750 ft, 10 (USNM). cottontail. Where the two occur to- Weston County: 4 mi E Four Corners, 1; 4 mi N, 3 mi W Newcastle, 1. gether, S. nuffaUii usually occurs at Additional records.— DAKOTA: SOUTH higher elevations in wooded or brushy Custer County: Custer (G. M. Allen, 1916: habitat, whereas S. auduhonii lives in 344); French Creek Cave, Custer State Park situations at lower (R. A. Martin, pers. com.); 4 mi W Fairhurn relatively open, grassy (band record: Tyers, 1963:17). Fall River elevations. For example, Merritt Gary County: 2 mi N, 1 mi W Burdock (band rec- obtained both species at Elk Mountain Cascade Cascade ord); (band record); Springs early in the 1900's; S. nuttaUii was taken ( band record ) . at 6000 feet, and S. auduhonii at 5000 feet. similar ORDER LAGOMORPHA—Hares, A ecological separation ap- exists between S. auduhonii and Rabbits, and Pikas parently S. jioridanus in the eastern section of the Family LEPORIDAE—Hares Black Hills (see account of S. jioridanus) . AND Rabbits Four females taken near Minnekahta Four species of lagomorphs, one of in mid-June 1967 were shot in arid grass- the genus Lepus and three of the genus land and pastures. Bluegrass, brome Sylvila', Valentine, Cherry Co., Ne- nuttallii. A female cottontail from 3 mi braska ) . E Custer Stebler reported by ( 1939:390) Sylvilagus floridanus occupies ripar- as Sylvi1a^ occur below 4200 feet in the individual the represents greatest pene- eastern section, where they are sjmpatric tration of this into the wooded species with, but ecologically distinct from, S. interior of the Black it Hflls; however, auduhonii. Sylvilagus floridanus has been was taken in an area that ". . . appears reported previously from the study area to be a natural for no evidence prairie, as Lepus sylvaticus by Bailey (1888: the of a former for- indicating presence 446), who noted that this rabbit is com- est was discovered" (loc. cit.). Sage- mon in bushy ravines in the northeastern bmsh, butter and Lin- goldenrods, eggs ( part of the Hills; however, specimens are eria and asters were vulgaris), conspicu- available only from Lawrence, Custer ous among various grasses at the site and Fall Ri\'cr counties. of capture. \\^hile a Ranger-Naturalist at Wind TURNER: MAMMALS OF THE BLACK HILLS 61

Cave National Park in tlic suninicr of gust (type locality. Black Hills, Hill City, Pennington Co., South Dakota). 1968, I had occasion to witness contrast- S. similis SylvUagus nuttallii grangeri—Nelson, 1909, N. ing habitat selection between /. Amer. Fauna, 29:204, 31 August. and S. a. Inulciji. Eastern cottontails are confined to brnshy draws, forest edges of Sylvilagus nuttallii grangeri, orig- \'alley systems that infringe into the foot- inally described from the Black Hills, liills, and thickets along sides of streams; occupies areas of sagebrush and timber desert cottontails frequent the open in the montane western United States. in prairie and upland grasslands. Several In the Hills, this species occurs spruce mornings were spent observing both spe- and pine forests of the boreal cap, down cies feeding within five feet of one through the transitional foothill zone to another on a closely mowed lawn near about 4500 feet; at the lower elevations Park Headquarters. To the north of the in the western section of the Hills, forest lawn was a brushy ravine of mountain edges and brushy draws are inhabited. mahogany, wild rose, hackberry {Celtis In such ecotonal areas, S. nuttallii is sym- occidentalis), and chokecherry; all else patric with S. audidjonii, from which it was upland prairie. When startled, S. often is difficult to distinguish (see ac- fioridanus sought refuge in the ravine, count of this species). Albeit the most whereas S. auduhonii behaved much as abundant lagomorph in the Black Hills, a jackrabbit and loped off across the specimens of Nuttall's cottontail are lack- grassland. At the north end of the ravine ing from Meade, Fall River, and Weston is Wind Cave proper; I found the skele- counties. therein on ton of an eastern cottontail I know of no instances of parapatry the rabbit 23 August 1967. Apparently, between S. nuttallii and S. fioridanus in the cave subse- had been washed into the study area, although such has been rain because bones were quent to a heavy reported from Laramie County, Wyoming the found about 80 feet below surface, (Hall and Kelson, 1951:53). Based on no or where there was vegetation signs the specimens examined, the nearest lo- of active inhabitation by this species. calities of record between these two lago- in the on 22 A female taken Park July morphs in the Black Hills proper are showed no of signs reproductive activity, approximately 15 miles distant [S. fiori- and a male obtained there on 31 August danus from Wind Cave National Park was a that was ex- juvenile parasitized (4100 feet), and S. nuttallii from Custer Ixodes Had- ternally by ticks, spinipaJpis (5300 feet)]. Whereas Nuttall's cotton- wen and Nuttall, and chiggers, Eutrom- tail inhabits wooded environs above 4500 bicula alfreddugesi (Oudemans). Two feet in the central and western Hills, the a taken near adults, a male and female, eastern cottontail occupies brushy ra- A. Hot Springs in April 1950 by J. King, vines and streamside thickets below 4200 were molting over the rump. feet in the eastern Hills and along the Specimens examined (6).—SOUTH DA- northern and southern peripheries. Both KOTA: Lawrence 7 mi N Spearfish, County: species are distributed outward from 1. Custer County: Wind Cave, Wind Cave the Hills along riparian communities on National Park, 1; Headquarters, Wind Cave the S. fioridanus to the east and National Park, 4100 ft, 2. Fall River County: plains, 2 mi N Hot Springs, 1 (UMMZ); 1 mi S Hot S. nuttallii to the west. Intergradation Springs, 1 (UMMZ). between nuttallii and fioridanus has been Additional record.—SOUTH. DAKOTA: suggested as possibly occurring along the Pennington County: Rapid City (Bailey, 1888: eastern base of the Rocky Mountains, but 446). Hall and Kelson (loo. cit.) concluded

Sylvilagus nuttallii grangeri (J. A. Allen) that this is not the case, partially on the Nuttall's Cottontael basis of a study of specimens from the Black HUls. Lepus sylvaticus grangeri J. A. Allen, 1893, from southeast of Hill Bull. Amer. Mus. Nat. Hist., 7:264, 21 Au- Individuals 62 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

1 Elk 2 City, Pennington County, were collected Custer, (UMMZ); Mountain, (1 USNM, 1 UMMZ); Redbird Canyon, Elk from pine-clad uplands and spnice-filled Mountain, 6000 ft, 1 (USNM). near an abundance of canyons, usually WYOMING: Crook County: De\ils Tower, rocks, fallen logs, shrubbery, or old 3 (USNM); Sundance, 1 (USNM). buildings, all of which are in close prox- townsendii Hollister imity to small open grassy areas. On 7 Lepus campanius July 1967, a melanistic juvenile which \\^HITE-TAn.ED JaCK RaBBIT had under a sought refuge ponderosa Lcpus campestris Bachman, 1837, J. Acad. Nat. pine log was captured on a wooded hill- Sci. Philadelphia, 7:349 (type locality, side near Nemo, Lawrence County. plains of Saskatchewan, probably near Carl-

ton House . A female obtained in Palmer Gulch, ) Lepus townsendii campanius—Hollister, 1915, Pennington County, on 12 April 1946 Proc. Biol. Soc. Washington, 28:70, 12 carried three that were 29 in embryos March, a renaming of Lepus campestris crown-rump length; another taken near Bachman. the same site on 13 August 1950 con- ^^ hite-tailed jack rabbits inhabit up- tained four embryos that were 17. A lac- land grasslands of the northern plains tating female was captured on 10 Au- and montane regions of western United gust 1929 at Elk Mountain, and a preg- States. Abundant on the surrounding nant female containing five embryos that flats, Lepus townsendii is less common 56 were ( crowni-rump ) was shot north- in the Black Hills proper. It inhabits east of Custer on 20 June 1967. Juveniles open woodlands up to 6000 feet and iso- and subadults of this species are common lated grassy areas such as Reynolds from mid-April to early September. Prairie, Gillette Prairie, and the Bald The obtained on 10 holotype, August Hills. Although no specimens are avail- 1894 W. W. is a full- by Granger, nearly able from Custer and Fall River counties, grown young of the year in subadult this species often has been observed ( post-juvenal ) pelage; whereas, four along roads at night in these areas. paratypes are juveniles in fresh juvenal Bailey (1888:438) recorded the first rep- and another is an adult with pelage, resentatives of L. townsendii from the worn pelage that was commencing to Hills (as L. campestris), and Palmer molt. New hairs can be observed under- (1897:28, 74) indicated that a single the older on the of a lying pelage rimip hunter near Newcastle, Weston County, obtained on 25 and topotype April 1910, killed more than 100 indi\iduals in on adults taken in and generally June 1893-94. early July. Apparently, adults molt to At lower elevations, these lagomorphs summer pelage from late March to mid- inhabit open prairie and upland grass- July, whereas to winter change pelage land. I have observed them in Wind begins early in August on some individ- Cave National Park and in pastures just uals. The melanistic is juvenile entirely southeast of Minnekahta; in such situa- black except for diffuse ochraceous guard tions L. tonnsendii o\erlaps ecologically hairs over the sides, venter, and white with SyIvilaL!,tis aiidu1)onii. At higher hairs along edges of the ears and on the elevations, L. townsendii frequents open tail. woodlands of aspen, pine, and spruce examined — Specimens (50). SOUTH DA- that border meadows and pasturt\s; un- KOTA: Lawrence 2 mi S County: Tinton, der these conditions, L. townsendii over- 6100 ft, 1; 2 mi W Nemo, 4700 ft, L Penning- with nuttaUii ton County: Hill City, 7 (1 USNM, 6 AMNH); laps ecologically Sylvikig,tis Redfem, 10 (USNM); Palmer Gulch, 3-4 mi to an undetermined extent. Hill SE City, 5300-5400 ft, 15 (9 UMMZ); A female shot southeast of Hill 16 mi NW 2 near Mount Custer, (UMMZ); Cit\' on 12 April 1946 contained four Rushmore, 1 (UMMZ). Custer County: Palmer embryos that were 70 in crown-rump Gulch, 8 mi SE Hill City, 1 (FMNH); 5.75 mi two males taken on 15 1965 N, 5.75 mi E Custer, 5220 ft, 1; Custer, length; June 3 (USNM); Bull Springs, 2 mi N, 9 mi W and 28 June 1967 had testes that were TURNER: MAMMALS OF THE BLACK IIILLS 63

49 and 52 in length, respectively. Yonng a di\ersitv of lilc lorms and habits. of the year were present from mid-March Among terrestrial squirrels, three are fos- to late August; in 1894 Palmer (1897:28) sorial (Cijnomys, Spermophilus, and observed parturition of individuals at Mannota) and one is scansorial {Eutarn- Newcastle, Wyoming, on 23 and 24 May. ias); two arboreal squirrels are scansorial Specimens obtained throughout the (Sciurus and Taniiasciurus) and one is colder months were in white pelage [V. volant (Glauconiys). Except for noc- field file turnal all other taxa are H. Cahalane ( notes, on at Wind flying squirrels, Cave National Park, p. 66) observed an diurnal and are among the most con- indi\idual in which the change to the spicuous members of the Black Hills white winter coat was well-ad\'anced on mammalian fauna. Seasonal activity 3 October 1934]. The presence of new varies for the different species, some hi- brownish hairs beneath the well-worn bernate in colder months, whereas others white hairs indicated that the pregnant remain active the year around. female mentioned above was undergo- minimus A. ing molt from winter to summer pelage. Eutamias pallidas (J. Allen) A male shot southwest of Sturgis was Least Chipmunk infested with ticks, Dermacentor ander- [Tamias quadrivitatus] var. palUdus J. A. Allen, soni Stiles. 1874, Proc. Boston Soc. Nat. Hist., 16:289, Specimens examined (17).—SOUTH DA- June (type locality restricted to Camp KOTA: Lawrence County: Deadwood, 1 Thorne, near Glendive, Dawson Co., Mon- (US\M). Meade County: Fort Meade, 1 tana, by Cary, Proc. Biol. Soc. Washington, (USNM); Black Hawk, 1 (NRW); Vanocker 19:88, 4 June 1906). 7 mi S, 1 mi W Stnrgis, 4800 ft, 1. Canyon, Eutamias lyiinimus pallidus—A. H. Howell, Pennington County: Rapid Cit>', 3 (USNM); 1922, Jour. Mamm., 3:183, 4 August. 3.5 mi i\, 1.5 mi E Deerfield, 1; Hill City, 1 5300 2 (UMMZ); 3 mi SE Hill City, ft, This pale chipmunk is widely distrib- (UMMZ); 3 mi W Rockerville, 1 (UMMZ). uted in the western section of the North- WYOMING: Crook County: Devils Tower, ern Great Plains, and inhabits riparian 1 (USNM); Sundance, 6000 ft, 1 (USNM). as well as shnab- Weston County: Newcastle, 3 (USNM). and scarp woodlands land habitats. In the Black Hills, E. m. Rodents ORDER RODENTIA— pallidus occurs in the forested or brushy lower southeastern foot- Rodents are by far the most numerous ravines of the of Black Hills mammals, both in number hills. Red Valley, and adjacent plains, at of kinds and in number of individuals. elevations up to 4300 feet. Specimens from Wind Cave National Park and from Seven families of rodents, totaling 22 southwest of Minnekahta are re- species of 18 genera are indigenous to clearly families ferrable to pallidus, whereas all other the study area; two ( represented examined from the Black by three genera, each of which contains chipmunks Hills are to the en- but one species) have been introduced proper assignable demic silvaticus the into the Hills. Four species of rodents race, (see following Distribution of E. minimus (representing two additional genera) account). one of two of two ha\'e been incorrectly reported from the represents examples of the same mammalian spe- region, and five species (representing subspecies cies within the defined bound- two additional genera) may possibly occurring Hills account of occur in the Black Hills. On the basis of aries of the Black (see and Nadler the numbers of individuals and resultant Dipodomys ordii). Sutton described two biomass, rodents are the base of the (1969:526) karyotypes (A and indi- vertebrate food-chain in the Hills. and B) in the genus Eutamias, cated that absence of heteromorphic Family SCIURIDAE—Squirrels chromosomes is evidence that the sub- Allies AND species characterized by these two dif- not Eu- Sciurids in the Black Hills represent ferent karyotypes do intergrade. 64 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

differs from E. m. silvoticus in tamias minimus pciUidus and E. m. sil- being paler over the and underside vaticus both possess karyotype B; speci- upper parts, sides, mens of intermediate characteristics are of tail. When compared with specimens encountered in the lower foothill zone, from eight other localities in the Hills, near Elk Mountain and Sundance for individuals from near Minnekahta and Wind Cave National Park are example. signifi- more in color tone see Table Little ecological data are available cantly pale ( re- Selected external and cranial meas- for E. m. pallidus in the Black Hills 8). abun- urements of three male and four female gion. Individuals are extremely from the Black Hills are: total dant along the slopes of Wind Cave pallidus Canyon, which is vegetated by mountain length, 203.0 ±5.66, 197.0 ±9.80; tail 75.3± hind mahogany, wild rose, hackberry, choke- length, 89.7±3.53, 12.53; foot ear cherry, wild plum, pricklypear cactus, length, 32.0±0.00, 32.0±0.82; Ca- pricklypoppy, and soapweed. V. H. length, 16.7±1.41, 18.4±1.89; weight, halane observed this small sciurid con- 52.9±3.96, 53.3±3.39; greatest length of suming dried wild cherries and wild skull, 32.7±0.18, 32.8±0.84; zygomatic file at Cave 18.4 18.2 brain- plums (field notes on Wind breadth, ±0.46, ±0.30; National Park). On 29 July 1968, a fe- case breadth, 15.9±0.14, 15.6±0.41; ros- male olive-backed pocket mouse cap- tral length, 11. 7 ±0.28, 11. 7 ±0.44; inter- tured in a snap-trap was partially orbital breadth, 7.5±0.00, 7.0±0.53; devoured by a chipmunk that departed maxillary tooth-row length, 5.2 ±0.04, as I approached. Perognathus jasciatus, 5.2±0.17; cranial depth, 13.3 ±0.14, 13.2 P. Jiispidiis, Pewmyscus manicidatus, ±0.48. Microtus Reithwdontomys megalotis, Specimens examined (15).—SOUTH DA- ochrogaster and M. pennsylvanictis were KOTA: Custer Countij: Beaver Creek Canyon, Cave National 4200 ft, 1; Elk taken in the same trapline with E. 7n. Wind Park, Mountain Campground, Wind Cave National pallidus in Wind Cave Canyon. Park, 4200 ft, 1; Headquarters, Wind Cave A male taken near Minnekahta in National Park, 4100 ft, 2; Wind Cave Canyon, over the mid-June 1968 was molting Wind Cave National Park, 4100 ft, 2; Wind rump and dorsum; scattered patches of Cave National Park, 4 (2 UMMZ, 2 WCNP). Fall River 0.5 mi 1.5 mi W Minne- new hair are interspersed among the County: S, kahta, 4200 ft, 5. older pelage on the sides and venter of another male obtained at the same time Eutamias minimus siK aticus White and place. Two adult females captured under these circumstances showed no Least Chipmunk

of molt still were in winter . sign ( pelage ) Eutamias minimus silvaticus White, 1952, Univ. Molt was manifest over the shoulders and Kansas Puhl., Mus. Nat. Hist, 5:259-262, dorsum of a female collected in Wind 10 April (type locality, 3 mi NW Sundance, 5900 Crook Cave National Park on 30 August 1968. ft, Co., Wyoming). Four placental scars were evident in The subspecies silvaticus is endemic the uterus of a female obtained on 13 to the Black Hills. Eutamias minimus is June 1968, but three other females col- represented by three morpliologically lected in the summer showed no signs similar, but distinctively colored, races of reproductive activity. The average on the Northern Great Plains as follows: length of the testes of five males collected the dark sdvaticus of the conifer-clad in the summer was 7.4 (7-9). Hills, the paler pallidus of surrounding Two males captured in Wind Cave foothills and wooded lowland escarp- National Park on 31 August and 1 Sep- ments and the extremely pallid caco- tember 1968, were parasitized externally demus of the barren alkaline topography by ticks, Ixodes spinipalpis Hadwen and to the east in the Badlands National Nuttall. Although external and cranial Monument and vicinity (Table 8). Least measurements are similar, E. m. pallidus chipmunks are extremely common and TURNER: MAMMALS OF THE BLACK HILLS 65

Table 8.—Geographic \arialion in coloration of the mid-dorsal pelage of adult Eutamias minivnis obtained in summer from the Northern Great Plains. Dashed lines separate populations that differ signifi- cantly.

o 0) c C « s Number and sex a! a O O of specimens o

150 (42 5, 108$ ) E. in. silvaticus. Black Hills of South Dakota and Wyoming Mean 28.4 46.4 28.8 24.8 SD ±3.33 ±0.42 ±0.27 ±0.28

12 (4ci,8$) E. m. paUidus, 2 mi N, 5 mi W Ludlow, Harding Co., South Dakota Mean 33.0 45.2 28.8 26.0 SD ±3.39 ±0.40 ±0.26 ±0.22

8 (65,25) E. 1)1. paUidus, 10 m S, 5 mi W Reva, Harding Co., South Dakota Mean 35.8 44.2 29.7 26.1 SD ±6.90 ±0.25 ±0.24 ±0.27

6(35,35) E. m. paUidus, northern Campbell County, Wyoming Mean 36.4 45.7 28.8 25.5 SD ±3.71 ±0.29 ±0.25 ±0.47

5(25,35) E. m. paUidus, 0.5 m S, 1.5 m W Minnekahta, Fall River Co., and Wind Cave National Park, Custer Co., South Dakota Mean 39.3 45.2 29.3 25.5 SD ±3.82 ±0.25 ±0.11 ±0.20

2 (15,15) E. m. paUidus, northern Siou.x County. Nebraska Mean 40.0 50.6 28.8 20.6 SD ±2.12 ±0.35 ±0.29 ±0.33

5(15,45) E. m. cacodernus, 14 m N Long Valley, Washabaugh Co., South Dakota Mean 58.0 43.7 30.1 26.2 SD ±4.05 ±0.25 ±0.16 ±0.13

4 (35,15) E. m. cacodernus, 10 m N, 4 mi E Potato Creek, Washabaugh Co., South Dakota Mean 69.9 44.4 30.5 25.1 SD ±9.42 ±0.24 ±0.14 ±0.10

6(15,55) E. m. cacodernus, 6 m S, 2 mi W Scenic, Pennington Co., South Dakota Mean 73.8 43.3 30.9 25.8 SD ±5.90 ±0.11 ±0.13 ±0.13

The widespread in all counties that comprise account of E. m. pallichis). species in the forested the Black Hills. Population densities is most numerous uplands seem to be much lower in the southern of the boreal cap and its presence has see been noted even near the summit of foothill zone and along the hogback ( 66 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Harney Peak (7240 feet) and atop Devils berries, asters, chokecherries, and various Tower. Eutamias minimus hibernates in grasses. A male shot in Ditch Creek the colder months. Individuals inhabit- Valley on 14 June 1965 was carrying a ing peripheral foothills and lower eleva- juniper berry in its mouth. Reforestation tions probably hibernate for a shorter programs in the Hills may suffer a 50 have been taken from to 75 annual seed loss due to period ( specimens percent 9 February to 20 November in these chipmunks and other small rodents areas) than do inhabitants of the colder (Henshaw, 1910:551); a single chipmunk recesses of the mountainous Central Ba- was observed visiting 38 seed spots in sin. Least chipmunks were first reported four minutes in one experimental plot, from the Black Hills by Baird (1858: which suffered 70 percent loss in a three- 299) under the name Tamias quadrivit- day period (Silver, 1924:167). Baits of tatus. oatmeal mixed with strychnine and wa- and wheat coated with hot tallow Individuals captured in the present ter, mixed with have study either were shot or were taken in strychnine previously been used to control museum special snap-traps baited with chipmunk popula- tions in the Black Hills loc. rolled oats. Eutamias minimus appar- (Henshaw, ently is ubiquitous, occurring in all habi- cit.). tats except extensive grasslands, which White (1953b: 588-589) described the are inhabited instead by Spermophilus process of molt in E. m. silvaticus, based tridecemlineatus. In areas where border- on study of a large series of specimens ing woodland interdigitates with grass- taken in several seasons of the year. I land, and in campgrounds, these two concur with his general analysis. Sum- small sciurids may be closely associated. mer molt of chipmunks in the Black Sparsely vegetated bluffs, rocky outcrops, Hills begins in mid-June and is com- and deteriorating abandoned buildings pleted in about mid-August. Seasonal are environs shared with Neotoma cine- charge in color affects mainly the tints rea. Pine-clad slopes, spruce-filled can- [tone and hue], whereas the overall pat- yons, deciduous riparian woodland and tern remains constant throughout the brushy meadows that contain piles of year, and is similar in both juvenal and logs are all inhabited by least chipmunks, adult pelages (J. A. Allen, 1890:49). which at one place or another have been Comparison of pelages of 53 adults taken in 37 females with obtained in association with Sorex cine- June ( 16 males and ) reiis, Peromijscus leucopus, P. manicu- 101 adults (28 males and 73 females) latus, Reithrodontomys megalotis, Cleth- obtained in July indicates that the new rionomijs gapperi, Microtus longicaudus, fur of late summer reflects reds at a sig- M. pemisylvajiicus and Zapus hudsonius. nificantly greater intensity than does the Eutamias minimus is most conspicuous older pelage of early summer. Measure- in old burned areas that are regenerating ments for the mid-dorsal region of speci- with secondary vegetative growth, and mens collected in June are followed by in logged areas with abundant trimmings those from July: color tone, 29.5 ±5.57, and fallen trees. For example, from 6 to 28.3 ±3.67; percent red reflectance, 44.7 8 September 1968, 76 individuals (39 ±0.43, 47.2 ±0.38; percent green reflect- males and 37 females) were obtained in ance, 29.7±0.25, 28.4±0.24; percent Roby Canyon along the western edge of blue reflectance, 25.6±0.27, 24.5±0.26. the Black Hills; these were taken mainly Change in pelage is manifest either by in logged areas near Roby Springs and distinct molt lines, or by interspersion on mountain mahogany-dominated slopes of new hairs beneath the older fur. Most near the mouth of the Canyon. juvenile and subadult chipmunks ob- tained in the warmer months were in Bailey (1888:437) noted that chip- of indi- munks in the Black Hills feed largely some stage pelage replacement, upon seeds of wild roses, wild rye, snow- cating a gradual process as individuals TURNER: MAMMALS OF THE BLACK HILLS 67 mature throughout the summer. Adults White (1953a:566) and aging criteria

at lo- also followed 1953b : 587-588 . taken on the same day the same White ( ) eahty may evidence different stages of There were no significant differences in molt, whereas others show no sign of dimensions of adults and old subadults, molt. Pelage replacement may be de- and both age groups were consolidated la\ed in breeding females (Howell, when making taxonomic comparisons. In 1929:28). Of 70 adult females obtained contrast to the findings of White (loc. in the summer that showed signs of re- cit.), females were significantly larger in production (embryos, placental scars, or total body length, weight, greatest length

, manifested no of of lactation ) 80 percent sign skull, zygomatic breadth, length of molt; the corresponding figure for 44 rostrum, and length of maxillary tooth- non-breeding adult females was 58 per- row (although absolute differences are cent. rather small for some of these means, Of 54 adult females examined in minor variation in a parameter that is June, 15 showed no signs of reproductive uniformly constant will result in statis- There- activity, 18 were lactating, 17 had an tically significant differences). average of 5.1 (3-8) placental scars in fore, sexes were treated separately when the uteri, and four pregnant individuals making comparisons; secondary sexual carried 4.5 (3-6) embryos that were 13.7 variation in coloration was not apparent. (7-27) in crown-rump length. Twenty- In the Black Hills, specimens from four males captured in this period had the Bear Lodge Mountains, and western testes that were 8.5 (4-17) in average areas in general, are more reddish than length. Of fifty-seven adult females ex- individuals from elsewhere; conversely, amined in July, 27 were non-breeding, reflection readings obtained with green 20 were lactating, 10 evinced 5.0 (3-7) and blue filters are higher for eastern placental scars, and none was pregnant. populations. For example, localities in The average testicular length of 17 males the Hills were consolidated into five collected in July was 6.1 (4-10). None of groups and resultant average percent re- eight adult females examined in August flectance of reds for each group follows: was pregnant and reproductive activity Bear Lodge Mountains, 50.0 ±0.26; was not evident in two of these; however, northwestern section, 48.0 ±0.32; south- two others were lactating and 5.5 (5-6) western section, 46.7 ±0.24; southeastern placental scars were present in the repro- section, 44.7±0.54; and northeastern sec- ductive tracts of the remaining four fe- tion, 42.9±0.34. Individual localities males of this series. The average testic- within these sections also follow this ular length of six males was 6.8 (6-8) in general trend. A distinct pattern of August. morphological variation within the Hills Two females obtained west of Nemo was not discernible. Selected external cranial measurements of 54 males on 29 June and 3 July 1967, and a male and 194.9 taken at Roby Springs on 7 September and 129 females are: total length, 201.0± tail 81.0± 1968 were parasitized externally by ticks, ±12.82, 12.22; length, Dermacentor andersom Stiles. Other 8.45, 83.3± 10.27; hind foot length, 31.0 least chipmunks captured at Roby ±1.61, 31.8±4.73; ear length, 16.3±1.29, 46.8±3.89, 55.0± Springs on the same date harbored 16.1±1.55; weight of 32.4 another kind of tick, Ixodes spinipalpis 6.97; greatest length skull, ±0.69, 18.4± Hadwen and Nuttall, lice, Hoplopleum 32.6±0.48; zygomatic breadth, 18.6 breadth of arboricola Kellogg and Ferris, fleas, 0.36, ±0.39; braincase, MonopsyUiis eumolpi (Rothschild), and 15.7 ±0.39, 15.7 ±0.34; rostral length, interorbital fur mites, Dermacanis hypudaei (Koch). 11.4±0.30, 11.5±0.28; breadth, 7.0±0.32, 7.0±0.27; maxillary In considering geographic variation tooth-row in E. minimus, cranial measurements length, 5.2±0.14, 5.3±0.15; were taken in the manner described by cranial depth, 13.2±0.27, 13.2 ±0.23. 68 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Specimens examined (612).—SOUTH DA- County: 1.5 mi E Buckhorn, 6150 ft, 19; 4 mi KOTA: Lawrence County: Spearfish, 3 E Four Corners, 6; 9 mi N, 1 mi E Newcastle, (SDSU); 3 mi S Speai-fish, 4200 ft, 1; White- 2; 6 mi N Newcastle, 1; Newcastle, 3 (USNM); wood, 2 (USD); Tinton, 2 mi N, 13 mi W SE Newcastle, 1 (USNM). Lead, 5900 ft, 30; 2 mi S Tinton, 6100 ft, 78; UNSPECIFIED LOCALITY. Black Hills, Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 4 (2 AMNH, 2 SDSU). 5800-6000 ft, 7; Little Spearfish Canyon, Sa- Additional records.—SOUTH DAKOTA: voy, 8 mi W Lead, 3 (USNM); 4 mi S, 7 mi Pennington County: Rapid Citv (Howell, 1929: W Cheyenne Crossing, 1; Deadwood, 19 57). WYOMING: Crook County: 2.5 mi N (USNM); Strawberry Hill Campground, 3 mi Sundance (Sutton and Nadler, 1969:526). S, 4 mi E Lead, 1 (SDMT); Dumont, 1 1 2 mi (USNM); Minnesota Ridge, (SDSU); Marmota flaviventris dacota (Merriam) W Nemo, 4700 ft, 22; Nemo, 4700 ft, 2 (1 SDSU); Blackfox Campground, 8 mi NW Yellow-bellied Marmot 2 Meade County: Vanocker Rochford, (NRW). Arctomys dacota Merriam, 1899, N. Amer. Canyon, 3 mi S, 1 mi W Sturgis, 4600 ft, 1; Fauna, 218, 30 October (type locality. Black Haven Dams, 6 mi S, 1.5 mi W Sturgis, 4600 Hills, Custer, Custer Co., South Dakota). ft, 5; 6 mi S, 1 mi W Sturgis, 4500 ft, 1. M[armota]. fUaviventer]. dacota—A. H. Howell, Pennington County: Rochford, 3 (UMMZ); 1914, Proc. Soc. Washington, 27:15, 2 Feb- South Canyon, 3-4 mi W Rapid City, 5 ruary. (AMNH); Dark Canyon, 2 mi S, 5 mi W [M.] flaviventris dacota—A. H. Rapid City, 3800 ft, 9 (8 AMNH); Beaver Howell, 1915, N. Amer. 7 Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, Fauna, 37:7, April. 50; 3 mi N, 9 mi W Deerfield, 6500 ft, 5; 3 Yellow-bellied marmots are distrib- mi N, 7 mi W Deerfield, 6900 ft, 2; Deerfield uted in montane areas of the western Lake, 6400 ft, 1; Castle Creek, 6500 ft, 2 (UMMZ); 2 mi S, 1 mi W Pactola, 1 (SDSU); United States, being confined to moun- 5 Sheridan Lake, 1 Redfern, (USNM); tains, foothills and rocky canyons, and (SDMT); Rockerville, 16 (2 NRW, 14 not occurring on the plains proper. Baird UMMZ); 3 mi S, 1 mi W Rocherville, 2; 20 first noted the occurrence of mi N Elk Mountain, 6000 ft, 3 (USNM); 4 mi (1858:344) NW Hill City, 2 (UMMZ); Ditch Creek, 14 this marmot in the Black Hills, as Arc- mi Hill 6400 near W Citv, ft, 29; Clendale, tomys flaviventer. The subspecies dacota Hill Hill City, 19 ( 15 4 FMNH); Cit>-, AMNH, is named from, and restricted to, the 4 (1 NRW, 3 USNM); 2 mi S, 13 mi W Hill Black Hills of South Dakota and Wvom- Citv, 6400 ft, 1; 4 mi S, 16 mi W Hill City, 1; 17 mi NW Custer, 1 (UMMZ); 16 mi NW ing, where it inhabits rocky hillsides, Custer, 53 (UMMZ); Palmer Gulch, 3-4 mi crevices in bluffs, rockpiles in meadows, SE Hill 5300-5400 20 City, ft, (UMMZ); or abides beneath deserted buildings. Horse Tliief Lake, 3 mi W Mount Rushmore Altitudinal distribution is from the bo- National Monument, 5100 ft, 1; unspecified lo- real of the Hills, down to about 4000 cality, 5 (UMMZ). Custer County: Palmer cap this Gulch, 8 mi SE Hill City, 5300 ft, 5 (FMNH); feet. Specimens of species have been 3.5 mi S, 0.5 mi W Keystone, 5000 ft, 15; 5.75 recorded from all counties that comprise mi N, 5.75 mi E Custer, 5200 ft, 22; Roby the study area. Yellow-bellied marmots Springs, 4 mi N, 22 mi W Custer, 5400 ft, 12; hibernate the colder months; the Roby Canyon, 2 mi N, 22 mi W Custer, 5200 during earliest and latest seasonal records from ft, 4; Bull Springs, 2 mi N, 9 mi W Custer, 6500 ft, 8 (UMMZ); 0.9 mi W Custer, 1 the Hills known to me are 12 April and 5300 13 2 (SDMT); Custer, ft, (4 AMNH, 22 August, respectively. UMMZ, 7 USNM); Bismark Lake Camp- Distribution of Marmota ground, 1 mi E Custer, 1 (SDMT); 4 mi SW flaviventris Custer, 3; Squaw [=Grace Coolidge] Creek, 1 in the Hills coincides with the occurrence (AMNH); Custer State Park, 21 (UMMZ); of rocky outcroppings or other high van- Campbell's Ranch, Elk Mountain, 4800 ft, 5 tage points from which these animals (1 UMMZ, 4 USNM); Mayo, 1 (USD); 3 mi can sun themselves and survey the sur- SW Pringle, 2; unspecified localitv, 5 (4 UMMZ, 1 USD). rounding countryside. These vantage WYOMING: Crook County: Devils Tower, points are closely associated with mon- 4 Warren Bear Moun- (USNM); Peak, Lodge tane meadows, pastures, and fields of tain, 6000 ft, 3 (USNM); 15 mi N Sundance, alfalfa and clover. Vernon Bailey found 5500 ft, 6; 15 mi ENE Sundance, 3825 ft, 1; flowers, leaves, and seeds of vari- 3 mi NW Sundance, 5900 ft, 17; 1 mi N Sun- green milk-vetch dance, 1; Sundance, 8 (USNM). Weston ous plants, including (As- TURNER: MAMMALS OF THE BLACK HILLS 69

truiialu.s hi.siilcatus) and stonecrop (Se- occasion, the colony sensed the approach (lui)i doiiiilasii) in the stomachs of mar- of a man while he was yet one-quarter mots from the Black Hills (Howell, of a mile distant. After a series of 1915:13). whistles from the adults, young animals Specimens from Ditch Creek Valley, retreated toward the burrow systems, Beaver Creek \'alley, and Little Spear- while the older marmots remained alert fish Canyon generally were taken in habi- until the man passed from sight. Graz- tats as described above. An adult female ing activity increased once again about and seven juveniles (four females and 16:30 hrs (MDT) and continued until three of males ) obtained southwest Key- near dusk. stone were shot at the entrance of an old Two pregnant females obtained south- lead-zinc mine and in adjacent wood- east of Hill City on 12 April 1946 con- piles. The mine was on a hillside that tained five and six embryos that were 16 was clad in pine and bur oak; well-used and 9 in crown-rump length, whereas a pathways were evident leading from the female from near Keystone (22 June mine to a valley meadow of bluegrass 1967) and another from Little Spearfish and clover below. Canyon (1 August 1967) evidenced and two A field party from The University of eight placental scars, respec- Kansas dug an adult female and six tively. Lactating females were taken on young males out of a den in Renter Can- 13 and 22 June 1967, and on 8 July 1965; marmots were from yon on 4 July 1947. The burrow was in young common early a hillside that faced north and the tunnel July through August. Testicular length of an adult male collected on 12 ran obliquely upward about 10 degrees June 1965 testes six from the horizontal for approximately 12 was 12, and of subadult males in 1967 feet, leveled off and ran at a right angle captured early August for another six feet; no nest was located. had an average length of 8.2 (7-10). The and underfur A graduate student from the Museum venter, feet, tail, of from near are a of Natural History, Paul B. Robertson, juveniles Keystone dull dark with a brown- or spent several days observing a colony of brown, pale hairs over the dorsum yellow-bellied marmots (five adults and beige-tipped guard and in contrast to the col- seven juveniles) at a locality 4 mi W sides, brightly ored reddish hairs of adults. Nemo, Lawrence County. The colony Change from subadult to adult and an- occupied an area of about 500 square pelage nual molt in adults occurs from mid- yards that comprised an old homestead. June of the old Deserted buildings, rock piles, and wood- through July. Replacement is most noticeable over the mid- piles were situated in a meadow and a pelage and pasture. Activity of the marmots began dorsum, sides, rump; replacement about 6:45 hrs (MDT) on sunny morn- of the mantle (a cloak of long hairs the shoulders and anterior third ings, with more restricted activity on covering of the is more difficult to ob- cloudy days. After an initial period of dorsum) serve but on occasion new hairs are vigilance, marmots spent most of the evident. The adult female from near morning grazing on the meadow. Move- ment of adults was much more conserva- Keystone seemed to have a mange-like disease and lacked hair over the tive than that of the young; while adults entirely mid-dorsum. remained alert, young romped freely, stopping often to preen or graze. When Marmota faviventris dacota differs raised from M. luteola of south-central feeding in tall grass, marmots up /. Wyo- on their haunches much more frequently ming (Converse, Albany, and Carbon was counties in the contrast in color than in short grass, and feeding ) , greater more sporadic. In late morning and between the mantle and mid-dorsal pel- early afternoon, grazing activity declined age (Table 9) and brighter color of the and sunning behavior increased. On one mantle of the former. Howell (1915:50) 70 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Table 9.—Geographic variation in coloration (mean and standard devia- tion) of the mantle and mid-dorsal pelage of adult Marmota flaviventris obtained in summer from the Northern Great Plains.

Number and sex of specimens s o averaged TURNER: MAMMALS OF THE BLACK HILLS 71

Canyon, 6 mi S, 1 mi W Sturgis, 4500 ft, 2. numbers of this species on the surround- Pennington County: Beaver Creek Valley, 4 ing plains, but saw none while in the mi N, 10.5 mi W Deerfield, 6400 ft, 2; Dark black Hills; S. tridecemlineatus was first Canyon, 2 mi S, 5 mi W Rapid City, 1 noted therein (AMNH); Sheridan Lake, 6 mi NNW Key- by Bailey (1893:45). stone, 1; 5.4 mi E Mount Rushmore National Ground sciuirrels often arc^ observed dur- 1 Ditch Creek Monument, (SDMT); Valley, ing the day along roadsides and in the 14 mi W Hill Citv, 6400 ft, 7; 1 Tigerville, parkways of campgrounds. Traps placed (USNM); Palmer Gulch, 3 mi SE Hill City, in these areas took five specimens in 5300 ft, 2 (UMMZ). Custer Counti/: Palmer in Gulch, 8 mi SE Hill Citv, 5400 ft, 1 "(FMNH); Ditch Creek Valley mid-June 1965;

3.3 mi S, 0.5 mi W Kevstone, 5000 ft, 8; another was shot as it ran from its bur- 13 6 Doran's Custer, (7 USNM, AMNH); row after water was poured into the Ranch, 3 mi E Custer, 1 (UMMZ). Fall River tunnel system. In Wind Cave National County: 5.5 mi E Minnekahta, 4000 ft, 1. taken on 27 WTOMING: Crook County: Renter Can- Park, two individuals were yon, 1 mi S Warren Peak, 6000 ft, 8; Bear August 1968 along a fence that served Lodge Mountains, 1 (USNM). Weston as a corral for bison. The enclosed area County: 0.5 mi E Buckhorn, 6100 ft, 1; 1.5 was closely grazed, whereas exterior to mi E Buckhorn, 6150 ft, 1. the fence, natural and UNSPECIFIED LOCALITY: Black Hills, prairie grasses A. 2 (1 USNM, 1 FMNH). forbs were about knee-high. J. King obtained several S. tridecemlineatus in Spermophilus tridecemlineatus pallidas early April in an upland meadow that individ- J. A. Allen was being grazed by horses; one ual was unidentified seeds in Thirteen-lined Ground Squirrel carrying its cheeks. Fifty percent of the stomach Spermophilus tridecem-lineatus var. pallidus contents of a female taken at Custer in J. A. Allen, 1874, Proc. Boston Soc. Nat. July 1893 was composed of grasshoppers Hist., 16:291, June (nomen nudum). and other insects, the remainder being Spermophilus tridecemlineatus var. pallidus seeds and matter J. A. Allen, 1877, in Coues and Allen, Bull. vegetable (Bailey, U. S. Geol. Surv. Territories, 11:872, Au- 1893). to the mouth gust (type locality restricted Reithrodontomys megalotis, Peromy- of the Yellowstone McKenzie Co., Ri\er, scus maniculatus, Microtus ochrogaster, North Dakota, by A. H. Howell, N. Amer. and Eutamias minimus often were taken Fauna, 56:122, footnote, 18 May 1938). in the same trapline with the thirteen- Spermophilus tridecemlineatus olivaceus J. A. Allen, 1895, Bull. Amer. Nat. Hist., 7:337, lined ground squirrel. Long (1965:581) 8 November (type locality. Black Hills, stated that he had seen Spermophilus Custer, Custer Co., South Dakota). tridecemlineatus and Eutamias minimus "so associated in Custer County, The thirteen-lined ground squirrel in- closely South Dakota, that the latter could be habits dry grasslands of the central chased into the burrows of the former." United States, and is locally common within the Black Hills in areas of short Four of eight females (for which are taken grass such as moderately grazed pastures, reproductive data available) mowed borders of roadways and camp- from 12 June to 1 July, were pregnant, 17 grounds, and upland meadows. This averaging 7 (3-9) embryos that were another spermophile ranges through the foothill (6-27) in crown-rump length; transition zone up to 6500 feet, but is individual was lactating. Testes of five more common below 5500 feet. Spermo- males obtained in the same period were philus tridecemlineatus hibernates in the 10.4 (5-16) in length. Juveniles first ap- and study area from mid-September until peared above ground in early July, mid-summer. early April; the earliest and latest sea- were common by sonal records as evidenced by the speci- Seven individuals obtained in mid- mens examined are 8 April and 7 Sep- June west of Hill City were molting con- tember, respectively. spicuously over both sides and the dor- National Grinnell (1875:82) observed large sum. A female from Wind Cave 72 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Park harbored several ticks, Ixodes sculp- I have compared a large series of tus Neumann, and fleas, Thrassis joins specimens from the Black Hills with

( Jordan ) . typical pallidus from northwestern

J. A. Allen (1895b: 337) described a (Harding County) and southwestern tridecem- Bennett South north- new subspecies, Spermophilus ( County ) Dakota, lineatus olwaceus from the Black Hills; ern and western Nebraska (Sioux, he indicated that it differed from the Cherry, Keya Paha, and Cheyenne coun- nearby pallidus "in its much darker ties), northeastern Wyoming (Campbell, ground color and the olivaceus creamy and Weston counties), and southeastern white tint of the light stripes and spots." Wyoming (Albany, Goshen, and Laramie

A. H. oliva- counties . Howell (1938:113) placed ) Morphologically, spcrmo- ceus in the synonomy of pallidus, stating: philes from the Black Hills fall within "The type series of olivaceus from Custer, the range of variation of the comparative South Dakota, lias been compared with material and do not differ significantly a large series of typical pallidus and is from pallidus in anv dimension (Table found to agree closely with it." Long 10). 1965:581 reinstated olivaceus as a sub- of ( ) Ground color and coloration stripes species of S. tridecemlineati4S, indicating and spots on individuals of S. tridecem- tliat in addition to the difference in col- lineatus vary greatly, and most possible oration noted by Allen, olivaceus also is color combinations are present when a characterized by shorter nasals, larger large series of specimens is examined. auditory bullae, and longer external ears On the average, mid-dorsal pelage of than pallidus. thirteen-lined ground squirrels from the

Table 10.—Geographic variation in selected external and cranial measurements of adult Spermophilus tridecemlincatus from the Northern Great Plains.

Number and sex of specimens averaged TURNER: MAMMALS OF THE BLACK HILLS •73

3800 Crowfeet Hills is dark in tone, high in reflectance Spearfish, ft, 1; [=Crow Peak?], Black Hills, 1 (AMNH). Meade County: Black of reds, and low in reflectance of blues Hawk, 1 (AMNH). Penninpton County: 0.5 However, these individuals (Table 11). mi N Silver City, 1 (SDMT); Rapid City, 2 readily fall within the range of variation (NRW); Beaver Creek Valley, 4 mi N, 10.5 6400 Ditch Creek Val- of populations from Bennett County and mi W Deerfield, ft, 1; ley, 14 mi W Hill City, 6400 ft, 6; 2 mi S, from parts of W'Noming, and do not differ 13 mi W Hill City, 1; Pactola, 1 (USNM); significantly from them. Thus, I must Castle Creek, 6500 ft, 2 (UMMZ); Diamond with Howell and S. t. oliva- concur place S Ranch, near Rapid City, 2 (UMMZ); Palmer 7 ceus in the synonomy of S. t. pallidiis. Gulch, 3-4 mi SE Hill City, 53-5400 ft, (UMMZ); unspeciBed locality, 1 (UMMZ). Significant secondary sexual variation Custer County: Palmer Gulch. 8 mi SE Hill was not e\'ident. males aver- Although City, 5300 ft, 4 (FMNH); Harney National aged somewhat larger in se\'eral cranial Forest, 10 (UMMZ); Custer, 25 (1 UMMZ, Bull dimensions, females were slightly larger 18 AMNH, 4 USNM, 2 FMNH); Springs, 2 mi N, 9 mi W Custer, 10 (USNM); Elk in total length and length of tail. Addi- Mountain, 1 (USNM); Buffalo Corral, Wind tional measurements (not listed on Ta- Cave National Park, 4300 ft, 2; Wind Cave ble 10) of 31 S. t. pallidus from the Black National Park, 2 (WCNP). Hills are: tail 75.3 ±7. 16; hind length, WYOMING: Crook County: Bear Lodge foot length, 31.5±1.40; weight, lU.Sib Mountains, 5200 ft, 1 (USNM); 15 mi N Sun- 22.73; and breadth, 11. 1± postorbital dance, 5500 ft, 3; 1.5 mi NW Sundance, 5000 0.57. ft, 6; 1 mi N Sundance, 1; Sundance, 5 Weston 1.5 mi E Buck- Specimens examined (102).—SOUTH DA- (USNM). County: KOTA: Lawrence Countij: L.5 mi S, 0.5 mi E horn, 6150 ft, 4; Newcastle, 1 (USNM).

Table 11.—Geographic variation in coloration of the mid-dorsal pelage of adult Spermophilus tridecemlineatus obtained in summer from the Northern Great Plains.

O o C c c a 03 a a Numlier and sex o O o of specimens fl o averaged H o o 111 PL, bc S! pm ^ S

21 (10c?, 112) Black Hills, South Dakota and Wyoming Mean 33.8 47.3 28.9 23.8 SD ±8.30 ±0.17 ±0.10 ±0.09

South Dakota 4 ( 1 ,5 , 3 9 ) Harding County, Mean 45.0 45.3 30.1 24.6 SD ±6.89 ±0.22 ±0.11 ±0.15

3 (U,29) Bennett County, South Dakota Mean 32.8 47.2 28.4 24.4 SD ±4.65 ±0.32 ±0.12 ±0.19

7(25,59) Northeastern Wyoming (see text) Mean 41.9 45.4 30.2 24.4 SD ±4.74 ±0.14 ±0.17 ±0.09

see text 11 (56,69) Southeastern Wyoming ( ) Mean 47.4 45.6 29.2 25.2 SD ±5.36 ±0.16 ±0.18 ±0.14

5(2

Cynomys ludovicianus ludovicianus part of the canyon floor and adjacent ter- races. In (Ord) 1950, King counted 614 bur- rows on an 11.7 acre of this Black-tailed Prairie Dog sample plot town (52.5 per acre). Wind Cave Na- Arctomys ludoviciana Ord, 1815, in Guthrie, A tional Park supports seven naturally es- new geogr., hist., coml. grammar . . ., tablished prairie towns and three Philadelphia, Amer. ed. 2, 2:292, descrip- dog new ones that were initiated in the north- tion on page 302 ( type locality restricted to the "Tower," sec. 10, T. 34 N, R. 10 W, westernmost corner of the Park in 1965. Bovd Univ. Kan- Co., Nebraska, by Jones, Cynomys ludovicianus occupies about sas' Publ., Mus. Nat. Hist., 16:140, 1 Oc- 655 acres, or three percent of the non- tober 1964). forested portion (21,449 acres) of the Cynomys hiclovicianus—Baird, 1858, Mammals, Park. Total inhabited . acreage in Repts. Expl. and Surw ., 8(1):331, by prairie 14 July. dogs increased by 90 acres from 1963 to 1967. The black-tailed prairie dog is a char- This is active the acteristic inhabitant of the short-grass species year around, decreases somewhat plains of central United States. F. V. although activity the colder months and on Hayden (1862:145) first noted this spe- during cloudy Both and Koford found that cies in the Hills region: "The largest one days. King males lose more than females [prairie dog town] I have seen is near weight the Black Hills, north of the Big Shey- during winter, possibly indicating that males enne river [Cheyenne River]. This vil- are more active at that time. Bur- rows are constructed on floors of can- lage, though sometimes interrupted by and on where high ridges or hills is connected, and yons adjacent terraces, fine-textured chernozems and chestnut covers an area of over fifty square miles." soils enable to excavate The Shirttail Canyon prairie dog town prairie dogs in the southwest corner of Wind Cave deep, continuous passages; at Devils National Park was studied extensivelv Tower, many burrows were located in fine sand of the Belle Fourche River by J. A. King (1955). C. B. Koford also studied the Shirttail Barriers to of ( 1958 ) Canyon floodplain. expansion towns seem to be unsuitable colony, as well as the prairie dog town in prairie dog Devils Tower National Monument. These soils of adjacent slopes, heavy cover, and terrain. two reports were consulted frequently rough in preparing the present account. Bailey Earthen mounds thrown up around (1888:440-441) described a prairie dog the entrances to burrow systems favor colony that occupied about 40 acres at a establishment of pioneer forbs such as place 2 mi E Rapid City. bigbract verbena (Verbena hracteata), The Devils Tower prairie dog town stickweed (Lappttla redowskii), dog- comprises about 40 acres between the weed {Dyssodia papposa), and false Belle Fourche River and the pine-clad pennyroyal (Hedeoma hispida). Koford hills of the National Monument. The analyzed the vegetation along a transect town is at least 75 years old; in June through the Shirttail Canyon colony and 1894, Vernon Bailey noted "a good sized found that the center of the town sup- dog town occupies a level section in the ported 67 percent buffalo grass, 24 per- valley at base of tower" (USBS files). cent tumblegrass (Schedonnardus panic- Counts made in 1947 and 1955 indicated ulatus), six percent blue grama, and that the population was fairly stable at three percent western wheat grass. In about 750 prairie dogs. In the area where the peripheral area, blue grama and west- Cynomys are fed by tourists, the number ern wheat grass increased in abundance of burrows reaches a maximum density and the town border was dominated by of 100 per acre. Japanese brome grass (Bromus japoni-

The Shirttail Canyon prairie dog cus) . W^ithin the confines of a town, town occupies about 75 acres, covering prairie dogs usually cut down tall plants TURNER: MAMMALS OF THE BLACK HILLS 75 that otherwise would obstruct a clear were dug out by badgers, and coyotes field of observation; examples of such are a fairly common sight in towns early plants are woolly mullein, Russian thistle in the morning. Although no actual (Salsola kali), snowberry (Symphori- sightings have occurred since 1953, char- carpos occidentaJis), silver sagebrush, acteristic trenches of the black-footed and brome grass. Other species sucli as ferret were found in the National Park snow-on-the-mountain (Euphorbia mar- in 1969. King reported that a prairie in the ginata), clammyweed (Polanisia graveo- dog was killed by a golden eagle left winter of I observed an lens ) and \'arious mustards are erect, 1955; eagle presumably due to a bitter taste, presence swoop down upon an adult prairie dog of thorns, a tough integument, or other in the summer of 1968, but the prairie of protective devices of the plants. In addi- dog escaped into its burrow. Remains tion to the numerous plant species listed a young C. ludoviciamis were found in Koford included of a horned owl that was by King ( 1955:8 and 9), pellets great scarlet globemallow, cutleaf nightshade roosting in Wind Cave Canyon in July (Solanum triforum), pepperweed (Le- 1968. pidiiim densifiorum) and lamb's-quarters Mating (occurs in February and as important foods of Cynomys liidovi- March and two to 10 young are born ciamis in the Black Hills region. in early April after a gestation period of King reported that the Shirttail Can- 30 to 35 days. The pups emerge from yon colony was divided into fairly stable under ground in mid-May. Evidently, social groups of several members, each only one litter is produced per year. King group occupying a particular territory found no evidence of reproduction of approximately one acre in size and among yearlings and noted that mor- adult male with a each consisting of an tality was high in the young during the few females and their young. Even first (36%) and second (22%) years. a mother defends the though prairie dog King noted that fleas (Opisocrostis area of her nest burrow, Koford thought hirsutus), mites ( Africholaelaps f^Ias- it was that this improbable aggressive- f!,owi), and ticks (Ixodes sp.) parasitize ness limits the number of families; for prairie dogs externally in Shirttail Can- 11 nest burrows were found in example, yon. an area of only 1.5 acres in Shirttail Specimens examined (29).—SOUTH DA- Canyon. KOTA: Pennington Cotmit/: 0.5 mi E Rapid Expansion of a dog town presumably City, 1 (SDSU); Rapid City, 11 (USNM). Custer McAdam's Ranch, 3 mi NW occurs when the population density in- County: Wind Cave, 1 (WCNP); Wind Cave National creases shortly after the pups emerge Park, 5 (2 UMMZ, 2 UW); Elk Mountain, from below noted that ground. King 5 (UMMZ); 1 mi NE Wind Cave, Wind Cave emergence of pups in May raised the National Park, 2 (WCNP). Weston average population density from four to WYOMING: County. Newcastle, 4 15 acre. In contrast to (USNM). prairie dogs per Additional records.—SOVTYi DAKOTA: the behavior of most rodents, adult Meade County: Stnrgis (USES files). Custer a from prairie dogs move into new area, County: Pringle (USES files); Horseshoe Eend, which the young are repulsed. The long- 12 miNE Elk Mountain (USES files). est movement recorded by King was 800 Sciurus rufiventer feet beyond the edge of the colony. niger Search for food may be another cause of E. Geoffroy St.-Hilaire colony expansion. Fox Squirrel In early July 1968, I obsei-ved an Sciurus fi. Geofi"roy St.-Hilaire, 1803, adult badger and two young excavating rufiventer Catalogue des mammiferes du Museum Na- a burrow in Wind Cave Na- prairie dog tional d'Histoire Naturelle, Paris, p. 176 Sev- tional Park earh' in the morning. (type locality restricted to Mississippi Val- eral burrows in Shirttail Canyon also ley, probably between southern Illinois and 76 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

central Tennessee, Ijy Osgood, Proc. Biol. Specimens examined (5).—SOUTH DA- Soc. Washington, 20:44, 18 April 1907). KOTA: Pennington County: Rapid Cit>', 3 (1 2 0.75 mi SW 1 Sciurus niger rufiventer—Osgood, 1907, Proc. SDMT, NRW); Rapid City, Biol. Soc. Washington, 20:44, 18 April. (SDSU). WYOMING: Crook County: Sand Creek, The fox squirrel is a eommon resident SBeulah, 1 (UW). of timbered areas and urban communi- Tamiasciurus hudsonicus dakotensis ties of eastern and central South Dakota, A. but has been reported in Wyoming only (J. Allen) from Cheyenne and Laramie (Long, Red Squirrel 1965:598) \ind from the Black Hills Sciurus hudsonicus dakotensis J. A. Allen, 1894, (Maxwell and Brown, 1968:155). The Bull. Amer. Mus. Nat. Hist., 6:325, 7 No- distribution of this species in western \'ember (type locality, Squaw [=Crace Coolidge] Creek, Custer South Da- South Dakota is essentially dendritic Co., kota). along the major river systems; for ex- Tamiasciurus luulsonicus dakotensis—Hayman ample, throughout the valley of the and Holt, 1940, in Ellerman, The families White River. In the the distribu- Hills, and genera of living rodents, British Mus., tion of Sciurus niger characteristically 1:346, 8 June. follows that of the bur oak and other In the northern plains region, Tamia- deciduous on elements, particularly sciurus hudsonicus inhabits montane for- that lead from stream many slopes up ests and several scattered conifer-clad beds in the northern Hills and the Bear escarpments. The species is commonest Mountains, and in wooded ra- Lodge on the boreal cap of the northern Black vines of the foothill zone. Hills where both white spruce and pon- Spread of the fox squirrel in Nebraska derosa pine are predominant; the former (Jones, 1964:146) and North Dakota conifer is scarce in the drier southern has been well doc- (Hibbard, 1957:527) woodlands, and accordingly, population umented. This species probably would densities of Tamiasciurus are much not occur in the area were it not study lower in these areas. The red squirrel, for the occurrence of gallery forests along or chickaree, is found in the Hills at river systems across the plains that elevations above 3800 feet, but is most served as avenues of dispersal and for numerous above 5200 feet. introduction by man. I know of no in- Hayden (1859:708) first reported T. troductions into the Black Hills proper, hudsonicus from the Black Hills, and but fox are known to have been squirrels J. A. Allen (1894a: 325) later described introduced in some areas dakotensis is adjacent an endemic subspecies ( see (northwestern Nebraska, for example; Hall and Kelson, 1959:400, map 257). see Jones, 1964). However, dakotensis is not restricted to Hayden (1856:79) indicated that the the Hills, but occurs also in several out- nothernmost limit of the fox squirrel in lying areas. For example, Vernon Bailey the plains region was at the mouth of captured a female at Belle Fourche, Running Water [Niobrara Ri\'cr] on the Butte Co., South Dakota, on 21 October later 1887. B. a Missouri, but ( 1859:708) noted that W. Evermann obtained male this species reached the limits of its at Edgemont, Fall River Co., South Da- range at the mouth of the White River. kota, on 7 October 1892, and V. H. Caha- I have observed these sciurids in resi- lane took a male 5 mi S Hcrmosa, Custer dential areas in Spearfish, Rapid City, Co., South Dakota, on 12 December 1934. and Hot Springs, and I have heard the J. A. Allen (1877:689) reported a speci- distinctive bark of fox squirrel in Beaver men from Bear Buttes, Meade Co., South Creek Canyon in Wind Cave National Dakota. W. J. Hoffman (1877:97) noted Park. Residents report this squirrel as that this species "sometimes is found in being quite common along wooded oak groxes eight miles west of the Post streams east and north of Spearfish. [Grand River Militaiy Post, Corson Co., TURNER: MAMMALS OF THE BLACK HILLS 77

South Dakota]," but this report is open limb (two to four inches in diameter) feet of the trunk. The to question. Swenk (1908:81) and Visher within two loosely (1914:88) noted the former distribution constructed nests were roughly oval in inches of red squirrels in nortliern Nebraska and shape, measuring 15 (13 to 17) Harding County, South Dakota, respec- in length, 11.3 (9 to 13) inches in width, 7.3 to inches in In- tively; however, Jones (1964:337-338) and (6 8) depth. did not admit Taniiosciums to the fauna teriors of the nests were shapeless, un- of Nebraska, and Andersen and Jones lined spaces, lacking a well-defined nest or sum- (1971:375) reported that the species evi- chamber. These were temporary winter nests dently does not now occur in Harding mer nests, but characteristic of Count)', although they did record a speci- are tightly constructed twigs, lichens, men from the Long Pine Hills in adjacent mosses and other plant materials, and Carter County, Montana, as dacotensis. frequently were observed among the in the Specimens of T. Jiudsonicus tentatively dense foliage of white spruce assigned to dakotensis have been re- northern Black Hills. ported from southeastern Montana and Materials used in construction of sum- the northern Laramie Mountains in Wy- mer nests were varied, consisting of news- oming (Hoffman and Jones, 1970:374, paper scraps (probably from a nearby Fig. 7). abandoned Civilian Conservation Corps Hills the shredded cloth and tufts of This sciurid is active in the Camp ) , bark, of lichens and dried year around. Although this species does fur, and particles member of not hibernate, it may undergo periods of graminoid plants. Another P. examined quiescence during prolonged inclement the field party, E. Marks, weather. A few individuals can be ob- two of the above nests and found 18 served in summer at most hours of the dipteran larvae, three dipteran pupae, unidentified dav, but maximal activity occurs between five flea larvae, two fleas, 7:00 and 9:30 hrs (MDT) and from one pseudoscorpian and eight collembo- 16:30 hrs to dusk. Even though popula- lans therein. tion densities vary from area to area, the Tamiasciurus in the Black Hills seems red squirrel is ubiquitous in wooded en- to utilize cavities of trees more frequently virons; pine-clad slopes, spruce-filled than outside nests, especially when rear- canyons, and oak-bordered drainage sys- ing young. On 27 June 1967, a male was tems all are inhabited. In the latter shot from a den tree in Vanocker Can- habitat, Tamiasciurus is ecologically sym- yon, Meade County. The hollow pine patric with Sciunis niger to an unknown tree was 30 feet in height and about 1.5 extent. The fox squirrel is far less com- feet in diameter; the nest entrance was mon and its distribution in the Hills 2.5 inches in diameter and located near closely approximates that of bur oak and the top of the tree. After the den tree other deciduous elements. was pushed over, it was split open for While with a Museum of Natural a better view of the nest. Within the a History field party in 1947, R. B. Finley hollow was not one, but sequential studied several external nests (located series of nests, graduated in age from rather than in a on branches of trees, bottom to top, comprising about 10 linear of red in Crook den cavity) squirrels feet of nesting material—fine dried He located three County, Wyoming. grasses and "old-man's beard" {Chionan- a 30"" ponderosa pine nest trees on slope in- thus virginicus). The series of nests with a southeastern exposure; the trees dicates usage of the tree cavity by one or were from eight to 17 inches in diameter, more squirrels over a long period of time. and 30 to 60 feet in height. Nests were Another nest was extracted from a hol- built 22.3 (18 to 25) feet above the low tree northeast of Custer on 24 June ground and either were placed adjacent 538 cubic inches of to the trunk, or rested upon a horizontal 1967; approximately 78 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

dried grasses and finely shredded bark ductively inactive; the average testicular constituted the nest material. length of 18 July-taken adult males was 16.6 Foods of red squirrels in the Hills (5-25). consist mainly of seeds of coniferous Utilizing the aging criteria of Layne 47 T. trees, but bracket fungi and acorns of (1954:251), juvenile and subadult bur oak also are consumed. A conifer hudsonicus taken (between 10 June and 15 cone is stripped of all its scales as the August) in the Hills were grouped into classes. of squirrels eat the ovules, dropping the age Time capture was gnawed core to the ground. These cut- then projected backward to estimate time of birth of these 10.6 tings generally accumulate beneath den individuals; ac- were born in late 29.8 or nest trees, forming large middens ( percent March, cumulations of refuse about a dwelling percent in early April, 44.7 percent in late 14.9 in place). Some middens that I have ex- April, and percent early May. amined were several feet in depth and The occurrence of pregnant females in late coincident with attainment of exceeded 10 feet in diameter. Bailey June ( maximal summer testicular in ( 1888:436) noted that red squirrels abide length males intimates that some adult females in rocky bluffs and store cones in clefts ) among the rocks; he also reported ex- in the Black Hills produce two litters, in tracting tvi^o or three bushels of cones one spring and the other in mid- from a hollow pine that contained a nest. summer, or else that some females breed In addition to cone scales and cores, pine later. and spruce needles, gnawed twigs and Layne (1954:232-235) described the bones, and other litter of food-remains seasonal change in pelage of Tamias- comprise the bulk of such middens. A sciuriis hiidsoiiicus loquax in central New maze of underground burrows and York; in general, my observations of molt blindly-ending pockets quite often pene- in dakotensis, although somewhat less trate these deposits. In the northern detailed, concur with his. Adults replace Hills, cores of white spruce cones are body pelage twice each year. Progress more predominant in refuse piles than of molt is gradual and somewhat diffuse, are those of ponderosa pine. Frequently, usually lacking distinct lines. Adults col- abandoned barns or feed-bins, and attics lected in late May, throughout June, and of deserted buildings are sites of vast in early July were in various stages of middens and food caches. pelage replacement. Several lactating Vernon Bailey obtained two lactating females still retained irregular patches of adult females southwest of Custer on 26 winter pelage on the rump in late June May 1894, and Ned Dearborn captured and early July. Two adult males cap- another at Redfern on 30 May 1910. Five tured at Sundance on 18 August 1913 of 16 females collected in June were were in full summer pelage, as were six nursing, and the uteri of three others individuals obtained in October. An adult evidenced 4 (3-5) placental scars; four male obtained at Jewel Cave Na- additional adult females showed no signs tional Monument on 20 November 1967 was in of autumnal of reproductive activity. Four pregnant process molt; new red squirrels (the only ones encountered hairs were manifest beneath the older in this study) taken between 20 and 28 pelage over the shoulders, sides and ven- ter. loc. cit. noted that hairs on June carried 5 (4-6) embryos that were Layne ( ) the tail of adults are 21.5 (6-42) in crown-rump length. The replaced but once average length of testes of 17 adult males annually, during the fall molt. captured in the period 1 to 15 June was Young red squirrels obtained in June 14.2 (8-22), whereas that of 18 males generally were in fine soft juvenal pelage, taken between 16 and 30 June was 17.3 whereas seven young captured in mid- Fifteen of 26 females obtained in fresh simi- (8-31). July were subadult pelage ( in July were lactating and 11 were repro- lar to adult summer pelage) and two TURNER: MAMMALS OF THE BLACK HILLS 79 others were undergoing pelage replace- Sundance are somewliat paler (27.4± ment over most of the body. 2.68), and reflect reds (54.6±0.15) more A male taken 2 mi W Nemo on 2 intensely and greens (25.3±0.10) less than lilack Hills Jul)" 1967 was parasitized externally by intensely typical repre- ticks, Dcnnacentor amlcrsoni Stil(\s, and sentatives; conversely, seven individuals another male from southwest of Sturgis (three males and four females) from harbored chiggers, Miy(itwni])icula eso- south of Tinton are comparatively dark ensis (Susa and Ogota). In addition, (25.6±1.75), and reflect reds (51.7± red squirrels throughout the Black Hills 0.17) less intensely and greens (26.5±: than do other Hills were infc\sted with unidentified fleas ( un- 0.08) more intensely fortunately, collected specimens of these squirrels.

. Tamiasciunis hudsonicus dakotensis ectoparasites were misplaced ) Comparison of 24 male and 31 female differs from T. h. haileiji, which occurs T. hudsonicus from the Black Hills re- west of the Black Hills, in being larger vealed no secondary sexual dimorphism in most external and cranial dimensions, in red in coloration or external and cranial and more reddish color ( reflecting measurements. Eight squirrels (four hues more intensely, Tables 12 and 13). males and four females) from north of It differs from T. h. fremonti, to the south-

Table 12.—Geographic variation in coloration of the mid-dorsal pelage of adult Tamiasciunis hudsonicus obtained in .summer from the Northern Great Plains.

o

55(245,31$)

Mean SD 6(25,4$) Mean SD 7 (35,45) Mean SD 27 (165,115)

Mean SD 4(25,25) Mean SD 10(45,65) Mean SD 10(65,45) Mean SD so MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Table 13.—Geographic variation in selected external and cranial measurements of adult Tamiasciunis hudsonicus from the Northern Great Plains.

Number and sex TURNER: MAMMALS OF THE BLACK HILLS 81

mi 1. in Table 13 of 55 Hills specimens are: W Minnekahta, 4200 ft, Unsj}ecified county: Black Hills, 3 (1 1 1 tail length, 137.6 ±9.44; liind foot length, NRW, SDSU, USNM). 52.0±4.00; and skull depth, 21.8+0.80. WYOMING: Crook County: Devils Tower, examined SOUTH DA- Specimens (295).— 4 (USNM); 15 mi N Sundance, 5500 ft, 1; Lawrence 6 KOTA: County: Spearfish, (5 15 mi ENE Sundance, 3825 ft, 7; 3 mi NW 1 3 mi SDSU, USNM); Big Spearfish Canyon, Sundance, 5900 ft, 16; Sundance, 2 (USNM); 4200 S Spearfish, ft, 1; Big Spearfish Canyon, unspecified locality, 1 (SDSU). mi 4800 White- 5 mi S, 2 E Spearfish, ft, 1; Additional records.—SOUTH DAKOTA: 4 Tinton, 2 mi N, 13 mi W wood, (USD); Lawrence County: Lead (Chambers, 1948:8). Lead, 6000 ft, 4; 2 mi S Tinton, 6100 ft, 22; Custer County: 16 mi SW Custer (USBS files). Little Canyon, 2 mi S, 10 mi W Lead, Spearfish WYOMING: Weston County: head Cheyenne 5800 ft, 6; Little Spearfish Canyon, Savoy, 1 River (J. A. Allen, 1877:689). (USNxM); Annie Creek, 6500 ft, 1 (USNM); 8 mi W Lead, 2 (SDSU); Deadwood, 6 Glaucomys sabrinus bangsi (Rhoads) (USNM); 4 mi SW Lead, 1 (NRW); Terry Peak, 1 (NRW); 1 mi S, 7 mi W Lake Rou- Northern Flying Squirrel baix, 1; 4 mi N, 1 mi W Nemo, 4900 ft, 1; Scittropterus alpinus bangsi Rlioads, 1897, Proc. 2 mi W Nemo, 4700 ft, 10; 1 mi E Nemo, 4700 Acad. Nat. Sci. Philadelphia, 49:321, 19 ft, 1. Meade County: Vanocker Canyon, 4 mi July (type locality, Raymond, Bear Lake S, 1 mi W Stmgis, 4600 ft, 1; Haven Dams, Co., Idaho). 6 mi S, 1.5 mi W Sturgis, 4600 ft, 8; 9 mi S, sabrinus —A. H. 1 mi W Sturgis, 4800 ft, 3. Pennington County: Glaucomys bangsi Howell, N. Amer. 13 Rochford, 6 (UMMZ); 6 mi N, 2 mi W Silver 1918, Fauna, 44:38, June. Beaver 4 mi 10.5 mi W Cit>', 1; Creek, N, Glaucomys sabrinus occurs in mon- 6400 ft, 17; Beaver Creek, 3 mi N, Deerfield, tane habitats of the western United 9 mi W Deerfield, 6500 ft, 2; 5 mi S, 1 mi W and in forest areas north- Pactola Lake, 1 (SDSU); Pactola Dam, 1 States, boreal of (SDSU); 10 mi W Rapid City, 1 (SDSU); ern North America. In the Black Hills,

South 4 mi 6 ( AMNH ) ; Canyon, W Rapid City, this squirrel is most abundant in the Rapid City, 1 (SDSU); Dark Canyon, 2 mi spruce forests of moist canyon systems S, 4 mi W Rapid City, 3 (AMNH); Deerfield on the boreal cap, but down to Lake, 6400 ft, 2; Deerfield, 1 (SDSU); Red- ranges about 4500 feet. are fern, 2 (USNM); 5 mi S, 1 mi W Pactola Specimens lacking Lake, 1 (SDSU); 1 mi W Rockerville, 1 for Meade, Custer, and Fall River coun- Creek, south of Rapid City, (NRW); Spring ties; however, residents report that this 2 (MHM); Ditch Creek, 14 mi VJ Hill City, species is a common mammal through- 6400 ft, 37; Glendale, 8 (7 AMNH, 1 FMNH); out the and is active the Hill City, 3 (USNM); 2 mi SW Medicine study area, year Mountain, 1 (SDSU); 3-4 mi SE Hill City, around. Grinnell (1875:81) first noted 16 Summit 5300-5400 ft, (UMMZ); Peak, flying squirrels in the Hills, under the 5500 ft, 1 (UMMZ); Horsethief Lake, 3 mi name Pteromys alpinus. W Mount Rushmore, 5100 ft, 1; near Mount are Rushmore, 1 (UMMZ); 1 mi S Keystone, 1 Northern flying squirrels some- (NRW); 20 mi N Elk Moimtain, 6000 ft, 1 what gregarious in the colder months. 17 mi NW Custer, 2 (UMMZ); (USNM); J. A. King captured 10 individuals in a 16 mi NW Custer, 3 (UMMZ); unspecified spruce-filled canyon southeast of Hill 4 (1 SDSU, 3 UMMZ). Custer locafity, between 11 December and 29 County: Palmer Gulch, 8 mi SE Hill City, City three of these were obtained .5.300 ft, 2 (FMNH); 3.5 mi S, 0.5 mi W Key- March; 5.75 mi E stone, 5000 ft, 7; 5.75 mi N, Custer, from the same tree. These specimens 5220 4 mi N, 22 mi W ft, 8; Roby Springs, were taken in rat traps that were at- 5400 ft, 6; 4 ini N, 14 mi W Custer, Custer, tached to trees about seven feet above 2; 4 mi N, 4.5 mi E Custer, 5400 ft, 1; Tepee level and baited with Canyon, 14 mi W Custer, 3 (USNM); Custer, ground peanuts. 7 (1 SDSU, 6 USNM); Harvey National For- South of Tinton, remains of a flying Custer State est, 3 mi E Custer, 2 (UMMZ); squirrel were found in the center of a Park, 5 (UMMZ); Squaw [=Grace Coolidge] midden of pine cones at the base of a 2 Cave Creek, 5 (3 AMNH, FMNH); Jewel dead tree. National Monument, 2.5 mi S, 12 mi W Custer, Most of the recently collected speci- 5400 ft, 1; 7 mi SW Custer, 1 (SDSU); Elk mens have been obtained on Mountain, 1 (USNM), unspecified locafity, 1 by tapping dead hollow trees and the (UMMZ). Fall River County: 0.5 mi S, 1.5 shooting squir- 82 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY rels as they exited from the den. Trees counties, in Wyoming; Long (1965:599) selected as nest sites usually are high on later concurred with King's assignment of in the Black Hills to slopes wooded in pine and scattered flying squirrels G. s. spruce and aspen. The average height bangsi. of five nest trees was 18.6 (15 to 25) Neither secondary sexual character- a half feet, and all were about one and istics nor age groups of adults (age was feet in diameter; openings to these nests determined in the manner of King, loc. to feet above were 12.8 (10 18) ground cit.) evidenced significant variation. In- surface. The oval-shaped entrance to dividuals from tlie Hills were compared one nest was 57 by 67 millimeters in with specimens from Park, Teton, and diameter; depth and width of the nest Sublette counties, in western Wyoming. cavity were 145 and 100 millimeters, re- Mid-dorsal pelage of animals from the nest com- spectively; the spherical was Black Hills is slightly darker in tone, and posed of fine grass and "old-man's beard." reflects reds more intensely and greens from Flying squirrels were obtained less intensely than those from western nests east of Buckliorn, Wyoming, on 18 Wyoming; however, tliese differences are adult female July 1951 (one lactating nonsignificant. Means and standard de- and four juveniles, of which two were viations of these color parameters are males and two were females), and west given for six adults (two males and four 1967 of Nemo, South Dakota, on 29 June females) from the Hills, followed by adult female and two (one lactating those for nine adults ( four males and five on juveniles, one male and one female), females) from the western Wyoming: 4 July 1967 (one adult male), and on color tone, 39.1±2.94, 39.5±4.78; per- 5 July 1967 (one lactating adult female cent red reflectance, 51.2±0.12, 50.0± and three juveniles, of which two were 0.28; percent green reflectance, 26. 1± males and one was a female). 0.07, 27.3±0.16; percent blue reflectance, 22.7±0.16. Reproductive data essentially are 22.7±0.08, lacking for this species in the Black External and cranial dimensions of individ- Hills except for the nest-taken representatives from the Hills examined adult uals mentioned above. Testes of an in the present study agree closely with male taken on 4 July were 11 in length. those of 12 adults measured by King are in evidence Young flying squirrels (loc. cit.). However, specimens from short through August. These have a fine the study area are significantly smaller Juvenal pelage that generally is darker than those from western Wyoming in than that of adults; the tail is blackish total length, tail length, ear length, great- rather than cinnamon. Glaucomys sa- est length of skull, and zygomatic Ijrinus from the Black Hills were para- breadth. Means and standard deviations sitized externally by an unidentified flea. of sLx adults (two males and four fe- A. H. Howell (1918:38) referred two males) from the Hills, followed by those specimens of G. sahrinus from the Bear of nine adults (four males and five fe- Lodge Mountains to the subspecies males) from western Wyoming, are: individ- canescens, indicating that these total length, 300.8±11.43, 329.4±9.69; uals in skull charac- approached bangsi tail length, 133.7±5.28, 150.0±4.08; ters and in a more vinaceous slightly hind foot length, 40.0±3.35, 41.3±1.11; coloring on the upperparts. When addi- ear length, 23.3±4.27, 27.3±1.11; tional specimens were made available weight, 149.8±23.48, 167.5+33.78; great- through his own field work. King (1951: est length of skiUl, 40.4±0.69, 41.6±0.81; 469) noted that representatives of Glau- zygomatic breadth, 24.3 ±0.25, 25.2± comys from the Black Hills differed sig- braincase 19.3±0.33, 19.5 nificantly from canescens of North Da- 0^50; breadth, 9.0±0.39, kota, but closely resembled three speci- ±0.22; postorbital breadth, mens of bangsi from Park and Fremont 9.0±0.50; interorbital breadth, 7.7± TURNER: MAMMALS OF THE BLACK HILLS 83

0.30, 8.0 ±0.63; rostral longtli, 13.5 ± Thomomys talpoides nebulosus V. Bailey 0.32, 13.9 ±0.59; length of maxillary Northern Pocket Gopher tooth-row, 8.1 ±0.14, 8.1 ±0.35; cranial Thomomys talpoides nebulosus V. Bailey, 1914, 17.6 ±0.51, 18.1 ±0.55; ratio of depth, Proc. Biol. Soc. Washington, 27:116, 10 cranial to breadth of depth braincase, July (type locality. Jack Boyden's Ranch, 0.92 (range: 0.91-0.95), 0.93 (range: Sand Creek Canyon, 15 mi NE Sundance, 3750 Crook 0.89-0.96). ft, Co., Wyoming). The smaller size, slightly different The northern pocket gopher is wide- coloration, and isolation of G. sahrinus spread in the Black Hills and common where local environmental factors are in the Black Hills ( about 245 miles west of populations in Park County, Wyom- favorable. The species inhabits montane warrant further before meadows and disturbed areas ing ) investigation parklands, an accurate assignment to a subspecies of secondary vegetative growth, and can be made. For the present time, 1 croplands, but is most abundant in heav- tentatively retain the subspecific name ily grazed valley pastureland. Specimens the Hills ])an(isi for the flying squirrels in the are available from throughout Black Hills. save from Fall River County, but the earthen mounds characteristic of gophers Specimens examined (31).—SOUTH DA- their there. Based on KOTA: Lawrence County: Little Spearfish evidence presence Canyon, Timon Campground, 1 (USNM); 2 specimens at hand, the altitudinal range mi S 6100 2 mi 4700 Tinton, ft, 1; W Nemo, of T. t. nebulosus, a subspecies endemic 7. Pennington County: 1 mi N Sheridan ft, to the Black Hills, extends from 3500 to Lake, 1 (USNM); Castle Creek, 6500 ft, 2 6500 feet. 146 individuals ex- (UMMZ); 4 mi SE Hill City, 53-5400 ft, 16 Among of the sex was (UMMZ); T. 1 N, R. 6 E, sec. 8, 1; 17 mi amined which known, oiily NW Custer, 1 (UMMZ); 16 mi NW Custer, 41 are males, suggesting that females 2 (UMMZ). may be more readily captured in traps. WYOMING: Weston 0.5 mi N, County: Thomomys talpoides was first reported 3 mi E Buckhorn, 6200 ft, 5. from the Hills by Bailey (1888:450). Additional records.—WYOMING: Crook The holotype, four allotypes, and County: middle fork Hay Creek, Bear Lodge from Sundance were taken Mountains (A. H. Howell, 1918:38). specimens by Vernon Bailey late in August 1913 in Family GEOMYIDAE— potato fields. Those obtained northeast in 1947 Pocket Gophers of Beulah, Wyoming, early July were captured in fields of alfalfa. Speci- This family of fossorial rodents has mens from Palmer Gulch were trapped undergone its entire evolutionary history in winter in a valley hayfield, between in North and Central America. Pocket a pine forest and a small stream. Four gophers, so named because of the fur- females were taken in August 1934 in a lined cheek pouches that open exter- montane meadow on Doran's Ranch, 3 are where nally on either side of the mouth, rep- mi E Custer by Stebler ( 1939:390) resented by but one species in the Black prominent plants were the short grasses, Hills; a second species {Geomijs hursar- but clumps of sagebrush, goldenrod, but- ius) has been reported erroneously from ter and eggs, and asters were conspicu- the area. Food habits of pocket gophers ous. and their extensive subterranean burrow Fifteen individuals from Nemo (late marked mounds of systems, usually by June and early July 1967) were trapped some earth near each entrance, present in a grazed pasture of bluegrass, clover, and economic problems to agricultural and scattered forbs. Three of eight go- thus control methods livestock interests; phers trapped east of Four Corners early such as extensive trapping or poisoning in July 1965 were taken in a burned area programs have been carried out by on a high ridge on which secondary re- of and ranchers in the Hills region. generation ponderosa pine many 84 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY kinds of forbs had occurred; the others and 12 in crown-rump length, respec- were captured in a pasture of bluegrass tively. Enlarged uteri, possibly indicat- and clover. No specimens have been ob- ing recent implantation were noted in tained in heavily forested areas. three females obtained in early July 1965 in two others were In early July 1965, a member of a Beaver Creek Valley; Placental scars and field party from the Museum of Natural lactating. (7, 3, 3) in three females taken in History, H. H. Genoways, studied burrow were evident dis- in 1967 on 29 systems of T. t. nebulosus and their Lawrence County June, tribution in Beaver Creek Valley, Penn- 3 July, and 29 July, respectively. Females located breed once from mid- ington County. Mounds were probably annually ( in dense, sodlike grasses where the soil March through mid-June) and produce was extremely dark, moist, and fairly three or four young; thus, the occurrence free of rocks, as well as in areas of dry of seven placental scars in late June is rocky soils that contained much organic of interest. Placental scars in pocket material. Burrows generally were lo- gophers generally remain two months cated at the edge of mounds. The fol- after parturition (Tryon, 1947:18) but lowing average measurements (in milli- the number of live-born young may be meters) are of 16 tunnel systems in smaller than the embryo count; addi- indicate valley soils: depth from opening of tional scars probably resorption 110- of rather than mound to bottom of tunnel, 170 ( some embryos production at this date. Hansen 250); width of tunnel, 65 (50-80); and of two litters early of about height of mound above ground level, 75 (1960:331) reported resorption all examined in (60-90). Burrows located in rocky soils four percent of embryos in Colorado. generally were much shallower, and sys- a study of T. talpoides tems excavated by subadults were at Young of the year were captured first most 80 millimeters below ground level in mid-June in the Black Hills and are in and were situated relatively deep fairly common in collections made in black soils. July; for example, 10 of 15 individuals 29 to 4 This species often is active in the obtained at Nemo from June July are five of six early afternoon. A female was obtained 1967 are juveniles, as 2 in Weston County on 7 July 1965 at 15:00 specimens taken at Beulah on July attain about three- hrs (MDT), shortly after traps were set, 1947. Subadults late and and I captured two juveniles about 16:00 quarters of adult size by July hrs (MDT) in snap-traps that were set early August, although younger individ- in a montane meadow southwest of uals are still taken occasionally. Two in the Spearfish on 7 August 1967. Although traps placed same burrow system most of the specimens examined were at Nemo took an adult female and a on 3 1967. taken in the summer months, J. A. King juvenile female July obtained individuals from November Bailey (1915:18) described the proc- 1945 through April 1946 in Pennington ess of molt in T. talpoides. Progress of County. Mounds and tunnel systems the crescentic shaped molt wave is from formed under the snow usually are evi- the rostrum, toward the rump. Young dent in spring, a further indication that taken in late June and early July were the northern pocket gopher is active the in a fine, juvcnal pelage; many subadults year around in the Hills. trapped late in July were in post-juvenal Testicular length of an adult male molt. Two simultaneous molt lines were captured on 4 July 1967 was 12. Only two of common occurrence on spring-taken pregnant females were observed, one adult specimens, and five individuals evi- taken on 3 July 1967 at Nemo, South dence three successive molt lines in Dakota, and the other collected on 4 progress. Long winter pelage remain on until mid- July 1965, near Four Corners, Wyoming; the rump of most specimens Merritt each carried four embryos that were 9 July. A female taken by Cary TURNER: MAMMALS OF THE BLACK HILLS 85 on Elk Mountain, on 19 November 1901 maxillary tooth-row than other popula- was undergoing molt to winter pelage, tions in the Hills. whereas three others obtained there TJioniomys talpoides nebulosus in the about this time were in fresh winter Black Hills resembles T. t. InilJattis of pelage. Four specimens from tlie iilaek the plains to the west, north and north- Hills ha\e patches of wliite hair on the east, but is significantly darker in tone shoulder or dorsum, and a female from and larger in most external and cranial Nemo has a white star-shaped area on dimensions (see Table 14). Specimens the forehead. from outlying areas of the Hills (Bear A female from Pennington County Lodge Mountains, Devils Tower, Four (2 July 1965) was parasitized externally Comers, \'icinity of Beulah and near by two kinds of lice, Geomyduecus tho- Buckliorn, in Wyoming, and Rapid City, monujus (McGregor), and a new species in South Dakota) are paler than typical currently being described by R. D. Price nebulosus, showing a tendency toward and associates, University of Minnesota. huUatus, but are best assigned to the Analysis of nongeographic variation former. Thomoimjs talpoides nebulosus in northern pocket gophers in the Black differs from T. t. pierreicolus of the plains Hills revealed no secondary sexual di- to the east and south in the same manner morphism in external or cranial dimen- as described above for T. t. bullatus, but sions, but tone of color of seven adult to a greater extent. Selected measure- males collected in the summer (2S.6± ments of seven adult males and 24 adult 4.38) is significantly paler than that of females from the Hills are: percent red 20 adult females (23.9±4.30). Because reflectance, 45.9 ±0.24; percent green re- of the topographic diversity in the moun- flectance, 27.6±0.13; percent blue reflect- tainous terrain, and due to the relatively ance, 26.5±0.20; body length, 226.3± sedentary nature of pocket gophers, lo- 9.34; tail length, 66.6±6.58; hind foot calized populations display great vari- length, 29.1 ±2.36; ear length, 6.8± ability in the Hills. For example, indi- 0.88; weight, 140.6 ±19.78; greatest viduals from Nemo are darker in tone, length of skull, 38.9±1.61; zygomatic reflect blues more intensely, and have breadth, 23.2 ±0.94; rostral length, 17.1 greater lengths of skull, rostrum, and ±1.09; interorbital constriction, 6.6±

Table 14. —Geographic variation in selected color, and external and cranial measurements of adult Thomoimjs talpoides from the Northern Great Plains.

Number and sex of specimens averaged 86 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

the South 0.34; length of maxillary tooth-rr)w, 7.9 tion along Dakota-Wyoming ±0.39; depth of skull, 15.3+O.SO. border. Specimens examined (158).—SOUTH DA- fasciatus KOTA: Lawrence County: Big Spearfish Can- Perognathus olivaceogriseus yon, 9 mi S, 3 mi W Spearfish, 5000 ft, 2; Little Swenk 2 mi S, 10 mi W Lead, 5800 Spearfish Canyon, Olive-backed Pocket Mouse ft, 2; 2 mi S Tinton, 6100 ft, 2; 2 mi N, 4 mi W Nemo, 5100 ft, 3; Nemo, 5700 ft, 15; un- Perognathus flavescens olivaceogriseus Swenk, locality, 1 (NRW). Meade County: specified 1940, Missouri Valley Fauna, 3:6, 5 June Fort Meade, 1 (USNM). Pennington County: (type locality. Little Bordeaux Creek, sec. Rapid City, 2 (USNM); Spring Creek, south 14, T. 33 N, R. 48 W, 3 mi E Chadron, of Rapid City, 2 (MHM); Beaver Creek, 4 Dawes Co., Nebraska). mi N, 10.5 mi W Deerfield, 6400 ft, 3; Beaver Perognathus fasciatus olivaceogriseus—Jones, Creek, 3 mi N, 9 mi W Deerfield, 6500 ft, 3; Univ. Kansas Mus. Nat. 20 mi N Elk Mountain, 2 (USNM); Ditch 1953, Publ., Hist., 5:520, 1 August. Creek Valley, 14 mi W Hill City, 6400 ft, 1; 2 mi S, 13 mi W Hill City, 1; Palmer Gulch, The olive-backed pocket mouse in- 3 mi SE Hill Cit>', 5300 ft, 14 (UMMZ); Tiger- Iiabits soils on the high plains ville, near Hill City, 2 (USNM); Redfern, 4 gravelly oc- (USNM); 16 mi NW Custer, 3 (UMMZ); peripheral to the Black Hills, and Harney National Forest, 3 mi E Custer, 4 casionally, individuals are taken in the 6 (UMMZ); unspecified locality, (UMMZ). foothill transition zone. The species has Custer County: Palmer Gulch, 8 mi SE Hill been captured only at four localities City, 5300 ft, 4 (FMNH); 3.5 mi S, 0.5 mi within the Hills of which two are in W Keystone, 5000 ft, 1; Bull Springs, 2 mi N, 9 mi W Custer, 6 (UMMZ); Custer, 5 (4 South Dakota, and two are in Wyoming. USNM, 1 AMNH); Mayo, 3 (USD); Camp- Altitudinal range as indicated by these bell's Ranch, Elk Mountain, 4800 ft, 1 specimens is up to 4800 feet. (USNM); Elk Mountain, 7 (USNM); 1 mi N Wind Cave, Wind Cave National Park, 1 Most available specimens of this spe- (WCNP); Wind Cave National Park, 1 cies were obtained in the Hills area in Beaver Creek, 1 (USNM). (USNM); the early part of the present century, WYOMING: Crook County: 1 mi E, 0.5 mi but thev first were reported from the N Beulah, 3550 ft, 14; Beulah, 3500 ft, 6; Bear area Merritt Caiy Lodge Mountains, 6200 ft, 6 (USNM); Devils by Jones (1953:522): Tower, 1 (USNM); 1.5 mi NW Sundance, captured five females in early October Rattlesnake 5000 ft, 4; Sundance, 2 (USNM); 1903 in Custer County; Vernon Bailey 6000 2 Sand 5 mi Creek, ft, (USNM); Creek, obtained a male near Newcastle, on the above mouth of canyon, 5 (USNM). Weston western border of the Hills, on 24 May County: 1.5 mi E Buckhorn, 61.50 ft, 2; 4 mi in 1913 found a mum- E Four Corners, 8; Newcastle, 5 (USNM). 1894, and August Additional record.—SOUTH DAKOTA: mified individual of unknown sex near Lawrence County: Deadwood (Bailey, 1888: Sundance, Wyoming. 450). I obtained two specimens while trap- in Wind Cave National Park in Family HETEROMYIDAE—Pocket ping a female was taken on 29 Mice axd Kaxgaroo Rats 1968; July (partially eaten in the trap by a chip- Three two of hetero- Both species ( genera ) munk) and a male on 28 August. are in the mammalian myids represented indi\'iduals were captured in snap-traps fauna of the Black a fourth Hills; species baited with rolled oats and placed in occurs nearby and its status in the Hills sparse grass at the junction of a fence region remains uncertain. All members row and a gravel road; the fence en- of this family possess external, fur-lined circled a sewage-settling pond at the cheek pouches, and to varying degrees head of Wind Cave Canyon. Micwtus are nocturnal, fossorial, and saltatorial. These rodents inhabit the interface of ochro

Tlie female taken in 196(S was neither and was trapped along a high fence that pregnant nor laetating; reprodnctive data separates the National Park from Guster are unavailable for those females cap- State Park. Vegetation, mainly grasses, tined by Gary. Testes of the adult male on the Wind Gave side of the fence was from Wind Ca\e Canyon were 3 in tall and dense. Due to overgrazing by length. bison in Guster State Park, vegetation was and short on that side of the Specimens examined (8).—SOUTH DA- sparse KOTA: Custer County: Campbell's Ranch, Elk fence. The male was trapped in an open 4800 5 Mountain, ft, (USNM); Wind Cave habitat of sparse vegetation, where the Canyon, Wind Cave National Park, 4100 ft, 1. soil was rocky, being similar to a gravel WTOMLNG: Crook County: Sundance, 1 Microtus ochro

examined 7 . — ING : October 1903 and N. R. Whitney ob- Specimens ( ) WYOM Weston County: Newcastle, 7 (USNM). tained a male south of Cascade Springs, Fall River County, on 8 February 1957. Family CASTORIDAE—Beavers Two other individuals were taken along the eastern periphery of the Hills, south The beaver is the largest rodent in of Rapid City, sometime between 1899 North America. Prior to the appearance and 1911 by Henry Behrens. D. G. of European man in the region, beaver Adolphson recently obtained skulls of were of great abundance and played a this species from barn owl pellets col- major role in luring early adventurers to lected 15 mi S Hot Springs ( Martin, the Black Hills. For example, expeditions 1971a: 4; 1971b). of the Verendrye brothers in 1743, the The female from Elk Mountain, taken American Fur Company in 1811, Jedc- on 10 October, evidenced molt on the diah Smith in 1823, and untold number dorsum; no other life history data are at of traders, trappers, and mountain men hand for the specimens examined. in the early 1800's mainly were in search of beaver and the Specitnem examined (5).—SOUTH DA- premium prices paid KOTA: Penniufiton County: Spring Creek, for beaver pelts at that time. south of Rapid City, 2 (MHM). Custer Morgan (1868:137) indicated that Coiintt/: Campbell's Ranch, Elk Mountain, beavers migrate each year, particularly 4800 ft, 2 (USNM). Fall River County: south an area overstocked. He of Cascade Springs, 1 (NRW). when becomes noted that an annual migration down ordii terrosus Hoffmeister Dipodomys the Missouri River usually occurred in Ord's Kangaroo Rat June, and wrote as follows: "A trapper whom I met on the Missouri River in Dipodomys ordii terrosus Hoffmeister, 1942, below Fort in Nebraska Proc. Biol Soc. Washington, 55:165, 31 De- 1862, Pierre, South informed cember ( type locality, Yellowstone River, [present day Dakota], 5 mi W Forsyth, 2750 ft, Rosebud Co., me that beavers were then (May 27)

Montana . ) coming down the river; that he saw them Habits of this heteromyid are similar dailv and had taken over fifty." Inci- to those of the preceding subspecies. dentally, Morgan (1868:209) gave a de- Vernon Bailey obtained three males and scription of the natural vegetation along four females near Newcastle, Wyoming the Missouri River as it appeared in the in late May 1894. When he examined 1860's, before modification by white man. kangaroo rats from the Northern Great As evidenced by numerous houses Plains region, Setzer (1949:524 and 543) and dams along most streams, beaver designated those specimens from Elk were abundant in the Black Hills at the Mountain the (in Black Hills proper) time of the Custer Expedition in 1874 as D. o. hiteohis and indicated that other (Grinnell, 1875:82; Ludlow, 1875:15; individuals from near Newcastle (on the O'Hara, 1929:252) and the Newton- western of the Hills were in- periphery ) Jenney Expedition in 1875 (Dodge, 1876: tergrades between D. o. hiteohis and 123). Increased colonization caused re- D. o. terrosus, but were assignable to the ductions in numbers of beavers, and latter. D. o. terrosus is discernibly Bailey (1888:441) reported that in the darker and larger than hiteohis except for autumn of 1887, only a few were "still a shorter tail and external ear pinna found in places far back from settle- (Long, 1965:618). ments" in the Black Hills region. Inadc- Further collecting is requisite along (juate game protection laws were ineffec- South the western border of the Hills to pro- tively enforced bv the state of vide critical data concerning the status Dakota between 1871 and 1889 (Elley, of these two subspecies in that region. 1945:5), and intensive trapping steadily Until further specimens are available, I reduced population hn els. According to follow Setzer. Harris and Aldous (1946:348), the spe- TURNER: MAMMALS OF THE BLACK HILLS 89 cics was nearly extinct in western South locality, Apple Creek, 7 mi E Bismarck, Burleigh Co., North Dakota). Dakota by 1927, but, reports of Seton (1929c: 453) and District Forester Allen The beaver is found on every major S. U. S. in l)ut it Peck ( Biological Survey files) creek and river South Dakota, indicated tliat more than two tlionsand is most numerous in the Black Hills beaver inhabited the Black Hills National where liundreds of beaver dams decorate Forest at this time. In 1927, a nonpoliti- the countryside (Gilliland, 1968b:5). cal form of Game Commission was estab- Morgan (1968:130) first reported this lished loc. cit. and ( Elley, ) , responsible species from the study area when he ob- management practices were initiated by served that the numerous dams on the South Dakota Department of Fishes, streams of the Black Plills were unusu- Game and Parks in the 1930's. Beaver ally small (in comparison to those he were restocked in the Hills from popu- studied around Lake Superior), ranging lated streams elsewhere in South Dakota. from 50 to 100 feet in length and two For example, due to the surplus of beaver to three feet in height. along the James, Vermillion, and Sioux Castor canadensis missouriensis dif- rivers in the southeastern part of the fers from C. c. concisor of south-central state in 1940, 300 of these large rodents Wyoming in its slightly paler underparts, were transplanted to the Hills (Wildlife relatively broader zygomata, more open Tips and Briefs, 1:14-15, 8 November orbit, narrower nasals, and in the shape 1940). Sale of 1196 beaver pelts netted of the jugal (Long, 1965:622-623). trappers in the Black Hills $35,285.00 in Some stocks of beaver have been in-

cif. . Hills areas 1945 ( Elley, he. ) Adequate protec- troduced into the Black from tion, in conjunction with the relatively outside of South Dakota. For instance, low prices now being paid for beaver three young from Yellowstone National pelts, have allowed the species to in- Park were planted along Squaw [^Grace crease in the Hills to the point that it Coolidge] Creek in Custer State Park in now is overusing the habitat. In recent 1914, and another familv of beaver was years, beaver have caused extensive dam- added in 1916 (South Dakota Conserva- age to white birch, aspen, poplar, willow, tion Digest, 36:3, 1969). Sixty beaver and Cottonwood trees along many drain- were transplanted to the Hills from an Conservation source in a on age systems ( Sonth Dakota unmentioned 1932; pair Digest, 24:12, October, 1957). Jim Creek and another pair along Cold Dams of this aquatic mammal regu- Springs Creek produced five and three late runoff by impounding water, thus young, respectively (reports of Regional aiding in the control of erosion and the Ranger, Black Hills National Forest, U. S. maintenance of an adequate water table. Biological Survey files). In 1947, five Sediment gradually accumulates behind white beaver were observed along the dams and many former beaver ponds Grizzly Creek, a pair of which was cap- eventually are transformed into mead- tured and placed in the Custer State ows, thereby increasing the stock carry- Park Zoo (American Fur Breeders, 21:62, ing capacity of the range. Berner December, 1948; Game News of Penn- (1953b:9) noted that the three to four sylvania, 19:16, March, 1949). thousand beaver in the Black Hills Na- From 1936 to 1944, 34 artificial sites, tional Forest, in combination with the consisting of small dams and temporary grazing activities of cattle, have been, bank lodges, were constnicted in the Hills Harris and and still are, converting many brushy northern Black ( Aldous, bottomlands into bluegrass pastureland. 1946:349). One hundred and thirty-six beaver were released at these artificial Castor canadensis missouriensis Bailey sites and 259 were released at natural Beaver sites. Members of the latter group tended to move on to other areas, whereas Castor canadensis missouriensis Bailey, 1919, beaver became established on 31 of the Jour. Mamm., 1:32, 28 November (type 90 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY artificial sites. Of 100 beavers tagged in habit the Black Hills. The status of and released in the period 1942 to 1944, three additional cricetids (two cricetines 14 were retaken (op. c/f.:350); time be- and one microtine) in the Hills region tween release and recapture averaged remains uncertain, and another micro- 11.9 (1 to 33) months. One individual tine has been incorrectly reported from was obtained at the site of release; and the area. the remaining 13 individuals moved an As part of their reforestation program, average of 9.1 (3 to 30) miles before the U. S. National Forest Service main- being recaptured. The average rate of tains numerous ponderosa pine planta- movement was 3.2 (0.3 to 14.0) miles tions throughout the Black Hills. These per month. In all, 1892 beaver were sites are prepared by windrowing, and trapped in 700 square miles of the north- the resulting dense grass and forb com- em Black Hills from 1936 to 1945; in a position that develops in these windrows five-day period in November 1944, 25 provides excellent habitat for small mam- beaver were captured near the head of mals. Periodically, populations of micro- the South Fork of Boxelder Creek (loc. tines in these areas increase to levels that cit.). are not controlled by predators and tre- Gilliland (1968b:5) noted that col- mendous damage to pine plantings re- onies in the Black Hills consist of one or sults. Such a population buildup of voles more families (one to 12 beavers) of all occurred in winter of 1968-1969, when an 870-acre area in the Dis- age groups (i. e., parents, kits, and year- Bearlodge trict suffered lings). Two to eight kits usually are 75 percent pine seedling born between late April and early June. mortality (Fred Wild, Range Conserva- tionist, com.) and the Di- Specimens examined (5).—SOUTH DA- pers. Taylor vide-Blacktail in the KOTA: Lawrence County: Tinton, 2 mi N, Creek plantation 13 mi W Lead, 6000 ft, 1; 2 mi S Tinton, 6100 same district was nearly a total loss ( John ft 1. 1..5 mi Fennington County: Rapid Creek, C. Windsor, District Ranger, pers. com.). W Rochford, 5300 ft, 1 (UMMZ); Spring Most damage occurred under the Creek, south of Rapid Cit>', 1 (MHM). Custer cover of snow; bark was removed from County: Palmer Gulch, 8 mi SE Hill City, 1 (FMNH). the root collar upward to the extent of the protective snow cover. Pine seedlings Family CRICETIDAE—Nath^e up to one inch in diameter frequently were gnawed completely through and Rats, Mice, and Voles other species such as aspen and oak were This includes a assem- family large killed by girdling. Subsequent surveys of terrestrial or fossorial blage partiallv of damaged areas yielded from two to rodents two subfamilies in representing 42 mice per 100 trap nights. For example, the Black Hills. The more primitive on the Dry Beaver Plantations, Penning- cricetines are nocturnal in habit are and ton County, 20 Microtus penn.sylvanicus, characterized cheekteeth that bear by two Clethrionomys p,apperi and two two rows of longitudinal tubercles. The Sorex cinereus were obtained in 50 trap Neotoma is an its teeth genus exception; nights (Fred ^^^ild, pers. com.). The are formed instead of oc- cuspless, being Forest Service plans to attempt control clusal lakes of dentine surrounded by measures on all areas sampled that enamel. The more advanced microtines yielded 15 or more rodents per 100 trap be active for short may periods through- nights (John C. Windsor, pers. com.). out any given 24-hour period, usually following well established runways, and Reithrodontomys megalotis dychei are characterized by rootless, prismatic J. A. Allen cheekteeth that lack enamel on occlusal Western Harvest Mouse surfaces. Reithrodontomys dychei J. A. Allen, 1S95, Bull. Three cricetine four genera ( species ) Amer. Mus. Nat. Hist., 7:120, 21 May (type and three microtine five genera ( species ) locality, Lawrence, Douglas Co., Kansas). TURNER: MAMMALS OF THE BLACK HILLS 91 — Reithwdontomijs megalotis dtjchei A. H. How- and along the interface of forested areas. ell, 1914, N. Amer. Fauna, 36:30, 5 June. in J. A. King obtained two females snap- Reithwdontomijs megalotis dijchei is traps placed under a log and beneath distributed throughout the Great Plains an oxerhanging creek bank in a spruce- of central United States, inhabiting filled canyon southeast of Hill City, and grassy environs, weedy borders of culti- he took 12 additional harvest mice in a vated tracts, thickets of shrubbery, and three-week period (28 December 1945 riparian communities. This mouse is ac- to 15 January 1946) from a gopher tive the year around, and in the Black mound located in an upland pasture in Hills occurs at elevations up to 6200 feet, the same general area. it is most abundant in the tran- although Only eight of 56 R. megalotis from sition zone below 5400 feet. The first the Black Hills area that are housed in specimens of the western harvest mouse the Museum of Natural History are adult in area were obtained taken the study by females, so reproductive data are meager. in B. Bailey at Spearfish early August In 1968, harvest mice in Wind Cave Na- R. was not 1927, but meii,alotis reported tional Park evidently bred from early the Hills until much from later, by Jones spring to late summer. A pregnant fe- and Mursaloglu (1961:25). male was live-trapped and released in on 27 and This species is fairly common along Beaver Creek Canyon June, female was taken there on 2 the prairie border and through the foot- a lactating one hills of the region. Nine individuals from July. Two pregnant females, carry- southwest of Minnekahta were trapped ing two embryos that were 19 in crown- with four in mid-June 1967 along fencerows of rump length and the other in bluegrass, brome grass, cord grass, west- embryos (10 crown-rump length), in the same on 26 era wheat grass, and scattered forbs, and were obtained canyon often were taken in runways of Microtus July. Two additional pregnant females, that were six ochrogaster. The mixed-grass prairie each carrying four embryos lowlands that are interspersed with in crown-rump length, were captured bushes and young trees in Wind Cave near the Park Headquarters on 30 Au- National Park also are inhabited, al- gust and 1 September. In other parts of though to a lesser extent, by harvest mice the Hills, young of the year were ap- in association with Etitamias minimus, parent in late May to late August. Testes males 7.2 Perognathiis fasciatus, P. hispidus, Pero- of 11 summer-taken were (6-8) myscus manicidattis, M. ochrogaster, and in length. Mils musciihis. In the Red Valley and Juvenal pelage of this mouse is short, foothills, moist draws and riparian com- fine and dark grayish-brown in color munities that support growths of choke- (Jones and Mursaloglu, 1961:14). Five cherry, wild plum, wild rose, hackberry, adults from Fall River County taken in and other thickets of shmbbery provide mid-June 1967 were undergoing the proc- adequate cover for harvest mice. For ess of molt as evidenced by new hairs example, along the brushy streamside over the anterior three-quarters of the liabitat of Beaver Creek Canyon, 15 R. m. sides and dorsum, and by the line of de- di/chei were live-trapped in late June marcation between fresh summer pelage 1968, along with 18 M. pennsylvanicus, and the worn winter pelage still present 25 P. nuinicuJatus, 10 Zapus hudsonius, on the rumps of these individuals. Other and one juvenile Neotoma cinerea. Five specimens obtained at this time either of these small rodents were collected were in fresh summer pelage or showed near Hot Springs on 4 September 1968, no signs of pelage replacement. Three on grassy ridges and in apple orchards individuals captured in Wind Cave Na- that flank the Fall River. tional Park late in August 1968 were o\'cr most At higher elevations, individuals were molting simultaneously parts to collected in erosional ditches, hayfields, of the body from the summer pelage 92 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY the longer and more dense winter pelage. 9.47; hind foot length, 16.6±0.S1; ear Harvest mice from the Black Hills length, 13.6±1.33; interorbital constric- are parasitized externally by ticks, Ixodes tion, 3.2±;0.13; length of incisive fora- spinipalpis Hadwen and Nuttall, fur- men, 4.4±0.15; depth of skull, 7.8±0.1S. mites, Dennacarus hypudaei (Koch), Specimens examined (96).—SOUTH DA- laelaptid mites, AndrolaeJaps fahren- KOTA: Meade County: Black Hawk, 1 holzi (Berlese) and Omithomjssus ha- (AMNH). Pennington Countt/: Rapid City, 8 22 mi Hill 6200 coti (Hirst), chiggers, Euschoengastia (SDMT); Moon, W Citv, ft, 1; 3-4 mi SE Hill City, 53-5400 ft, 22 fasolla Brennan and Beck, fleas, Ma- (UMMZ); Spring Creek, 4 mi ENE Rocker- laraeus telchinum (Rothschild) and ville, 3600 ft, 1 (UMMZ). Custer County: an MonopsyUiis wagneri (Baker), and Roby Canyon, 2 mi N, 22 mi W Custer, 5200 unidentified Anopluran louse that is rep- ft, 2; 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; 3 mi N 1 5 mi 2 mi resented in collections only by the nymph Pringle, (UMMZ); N, E Wind Cave, Wind Cave National Park, 4200 stage. ft, 1; 1.5 mi N, 0.5 mi W Wind Cave, Wind of in col- The preponderance males Cave National Park, .3250 ft, 1; Beaver Creek lections from the Black Hills invalidates Canyon, Wind Cave National Park, 4200 ft, 13; Wind Cave National 4100 any attempts to test for secondary sexual Headquarters, Park, ft, 5; Wind Cave Canyon, Wind Ca\'e National dimorphism in R. megalotis. When study- Park, 4100 ft, 12; Wind Cax'e National Park, harvest mice on the Great ing Plains, 5 (UMMZ); unspecified locality, 1 (UMMZ). Jones and Mursaloglu (1961:12-13) Fall River County: 5.5 mi E Minnekahta, 4000 found that the females are, on the aver- ft, 1; 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 9; 1 mi N, 5.5 mi E Hot Springs, 3400 ft, 2; age, larger than males in all external and 1 mi N, 6 mi E Hot Springs, 3400 ft, 3; Hot several cranial dimensions. However, in- Springs, 1 (UMMZ); 4 mi E Hot Springs, 3400 dividual variation exceeded sec- greatly ft, 1; 2 mi S, 2 mi E Hot Springs, 3600 ft, 1. sexual variation and ondary statistically WYOMING: Crook County: 1.5 mi NW differences were significant lacking. A Sundance, 5000 ft, 3. of variation in coloration or size pattern Additional records.—SOUTH DAKOTA: in the Hills was not readily discernible Lawrence County: Spearfish (BB). in the present study. The remarkable uniformity in pelage Peromyscus leucopus aridulus Osgood color of R. over a considerable megalotis White-footed Mouse geographic range also was noted by aridulus Jones and Mursaloglu (1961:16). Most Peromyscus leucopus Osgood, 1909, N. Amer. Fauna, 28:122, 17 April (type grassland cricetine species on the Great locality, Fort Custer, Big Horn Co., Mon- Plains have a dark eastern population tana). and a pale western population. Environ- The white-footed mouse occurs mental conditions along riparian habitats the northeastern two-thirds across the plains are somewhat similar to throughout of North America. The distribution of conditions of more eastern habitats, pos- across the arid sibly lessening differential selection pres- Feromyscus leucopus Great Plains the sures of R. megalotis in comparison to closely approximates dendritic of mesic deciduous that incurred by nonriparian grassland pattern communities that follow major drainage cricetines (loc. cit.). No significant geo- The is un- graphic variation in color or size is evi- systems. species relatively common in the Black Hills. It inhabits dent in specimens examined from areas the discontinuous deciduous surrounding the Hills in South Dakota notably woodland formations to 6100 and Wyoming (Table 15). Cranial meas- up feet, urements were taken in the manner de- although it is more abundant below 5000 feet. No are at hand from scribed by Hooper (1952:9-11). Addi- specimens tional external and cranial measurements Fall Bivcr County, but white-footed mice (not included on Table 15) of 16 adults probably occur there. Although this from the Hills are: tail length, 62.5± mouse has been taken in the Hills only TURNER: MAMMALS OK 11 IK BLACK HILLS 93

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lO 94 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY from mid-June to mid-August, it is active of Spring Creek. This site is mentioned the year around and does not hibernate. as a further example of the restrictt^d first habitat of P. the J. A. Allen (1894a: 325) reported leucopus on plains. white-footed mice from the Hills under Spring Creek was "bordered by box- the name Sitomys americamis arcticus elders, cottonwoods, plum thickets, wil- (see below). lows, wild currants, and rank weeds and extends from at the time of visit loc. Big Spearfish Canyon grass" Granger's (

elevation is cit. . Savoy, where the streambed ) about 5000 feet, northward, emerging Throughout its range in the Black from the Black Hills at Spearfish (3600 Hills, the white-footed mouse is sym- Creek courses below feet). Spearfish patric with the deer mouse. These two massive with luxuriant clifi^s, vegetation species are often taken side-by-side in bur American boxelder, ( oak, elm, green the same trapline (see account of Pero- and deciduous on the ash, shrubbery) myscus maniculatus) and are frequently floor. Sixteen 11 males and five canyon ( difficult to difi^erentiate. For example, P. were taken in the females) leucopus J. A. Allen (1895a: 268) listed 39 speci- underbrush of this woodland in riparian mens of Peromyscus leucopus from Cus- early August 1967. Vanocker Canyon ter and 20 from Squaw Creek. He noted arises on the southeast of Dead- slopes that a few specimens presented a decided man Mountain (4800 feet) and extends fulvous or reddish wash, and that one at 3600 northeastward, Sturgis ( the emerging specimen ( AMNH 9370) approached feet). Five individuals (four males and characteristic "fulvous tint of nehracen- one were obtained in this can- female) sis" [P. maniculatus], Allen later (1899: on 28 and two more one yon June 1967, ( 15) assigned all of these specimens to a of each were collected there on 15 sex) new subspecies, '^Peromyscus texanus 1967. These mice were August trapped suharcticus [P. m. artemisiae]," stating in brush in a that bordered an pasture "Black Hills specimens are not at all and in a mixed aspen grove, oak, pine, typical, some of them decidedly ap- and woodland. aspen proaching P. t. nehracensis, while others five males Nine Pewmysciis leucopus ( are quite like the Montana and Saskatch- and four females) were captured in the ewan examples." Only 10 of the 59 speci- Central Basin, northeast of Custer, in mens listed by Allen are actually P. leu- late June 1967. These were taken in copus, the remainder being maniculatus aspen woodlands bordering Iron and (including AMNH 9370). Externally, Grizzly creeks; the woodland contained white-footed mice are larger in body much undergrowth, fallen logs, decid- size, have longer tails and hind feet, and uous shrubbery and rock outcroppings. the summer pelage is somewhat duller, Eutamias minimus, Peromyscus manicu- with a less distinctly bicolored tail, in latus, Clethrionomys gapperi, Microtus comparison with deer mice. Subadults longicaudus, and M. pennsylvanicus are of the former species are easily confused associates of P. in the Hills. leucopus externally with adults of the latter, but White-footed mice also inhabit allu- the larger, more robust skull of P. leu- vial wooded bottomlands of in drainages copus, coupled with age determination the Red and foothills. Valley Specimens by degree of tooth wear, readily resolves collected Creek in late along Squaw July this dilemma. 1894 by W. W. Granger were taken in Three of six adult female P. leucopus an area that, "is wooded with aspen, from the Hills, for which willows, boxelders and other deciduous reproductive data are available, showed signs of repro- trees" (J. A. Allen, 1895a: 261). Granger ductive One taken northeast of also captured eight P. leucopus well out activity. on the prairie east of the Black Hills, Custer on 26 June 1967 carried six em- about seven miles southwest of the mouth bryos that were 3 in crown-rump length. TURNER: MAMMALS OF THE BLACK HILLS 95

Another, obtained southwest of Sturgis Hills revealed no apparent secondary on 28 June 1967, was lactating and had sexual variation in coloration or in ex- five placental scars present in the repro- ternal and cranial measurements, nor ductive tract. The third, captured south was a pattern of variation in these param- of Sturgis on 15 August 1967, had four eters discernible in tlie Hills populations. placental scars. The average length of Specimens from the Black Hills, and testes of 11 adult males collected in the adjacent areas to the west in Crook summer was 11.8 (10-13). County, Wyoming, are darker in average Many adult white-footed mice taken tone of color, and much smaller in most in late June and early July were molting external and cranial dimensions when over the dorsum and sides, whereas compared to populations on the South other individuals already were in fresh Dakotan plains (Tables 16 and 17). The summer pelage. Juveniles have soft gray small size of the Black Hills representa- fur. Two subadults from northeast of tives has been a source of past mis- 19 1967 were identifications. Custer ( June ) undergoing in which new brown post-juvenal molt Feromijscus leucopiis probably ex- hairs were emerging along the sides, tended its range westward on the Great still furred in whereas the dorsum was Plains along major drainage systems in the grayish juvenal pelage. mesic Post-Wisconsin times. The pres- Examination of 26 P. leucopiis (16 ently isolated segments formerly were males and 10 females) from the Black part of a more or less continuous and

Table 16.—Geographical variation in coloration of the mid-dorsal pelage of adult Pewmyscus leucopiis obtained in summer from the Northern Great Plains.

Number and sex of specimens averaged 96 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Table 17.—Geographic variation in selected external and cranial measurements of adult Peromijscus leucopus from the Northern Great Plains.

Number and sex j- of specimens B m averaged [5 Ji TURNER: MAMMALS OF THE BLACK HILLS 97

species. Although it is most common in ods were not used in the current study, grassy cn\iions of the Red Valley and calculations of population size were not foothills, P. inaniciilatus occurs in a wide attempted, but, it is certain that Dice's variety of habitats, including cultivated computations underestimated the num- cropland, grazed or ungrazed pasture- ber of deer mice in the Black Hills. land, grassland meadows, marsh and In the northern foothills and canyons, stream borders, spruce and aspen wood- and along some drainages in the central lands, dense pine forests, and even section of the Black Hills, P. m. nehras- among bare rock at the summit of Harney censis is ecologically sympatric with P. Peak (7240 feet), atop Devils Tower, leucopus (see account of that species). and several hundred feet below ground I have taken both species within a few surface in Wind Cave. feet of each other, and even in the same Dice (1939:5) estimated that the to- trap, in Little Spearfish Canyon. How- tal population of P. maniculatus in the ever, P. leucopus is less common and is Black Hills in late summer of 1935 was stenoecious, generally associating with between one and five million individuals. elements of the eastern deciduous forest. He indicated that deer mice occurred in Due to its occurrence in all available the deer is associ- the greatest abundance in dense and habitats, mouse closely ated with each of the other terrestrial medium dense stands of western yellow mammals in the Hills [ponderosa] pine in densities of 932 and region. 894 mice per square mile, respectively Deer mice captured in Wind Cave {op. cit.:2. Table 1). However, Dice National Park in summer of 1968 were

1939 : 1 based his estimates on 39 into classes tooth ( ) only grouped age based on specimens and erroneously assumed that wear, general dimensions, and type of the Black Hills population was isolated pelage. Of 315 individuals, there were to a large extent. Due to poor trapping 188 adults, 116 subadults, and 11 juve- results in grasslands near Rapid City and niles. Mice taken in July (97) and Au- south of Hot Springs, Dice did not trap gust (203) were further segregated by in grassland, pasture, or cultivated fields sex. In the following analysis of popu- in other parts of the Hills (op. cit.:3) as- lation composition, percentages for July suming that no deer mice occurred in are followed by those for August: adult these areas (op. cit.A). males, 32.0, 32.0; adult females, 25.8, subadult subadult My own investigations indicate that, 28.1; males, 19.6, 12.8; although deer mice are omnipresent in females, 20.6, 22.7; juvenile males, 1.0, The the Black Hills, the highest population 3.4; juvenile females, 1.0, 1.0; analy- densities are attained in those environs sis indicated that about 40 percent of summer of P. maniculatus in dominated by graminoid vegetation. For populations in a 13-18 the Black Hills are immature individuals. example, five-day period ( June 1967) 82 deer mice were captured This species may breed most months near Minnekahta in arid pastureland of of the year in the Hills region, but ade- the data are available for the Red Valley ( see account of Microtus quate only

. A. ochwgaster for a description of the area) summer. J. King captured two preg- In approximately 2000 trap-nights in the nant females southeast of Hill City, one five summer of 1968, 315 P. nuiniculatus of which carried embryos ( 14 March were taken in the grassy uplands and 1946) measuring three in crown-rump semi-grassy canyon systems of Wind length, and the other containing two Cave National Park. On 27 August alone, embryos (25 March 1946) measuring 23 66 deer mice were captured in 150 trap- in crown-rump length. On 5 April 1946, nights. Most of these specimens were King obtained two gra\'id females north discarded after being examined for ecto- of Newcastle that each carried five em- parasites and autopsied for reproductive bryos which had an average crown- data. Because quadrat sampling meth- rump length of 2 and 3, respectively. 98 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Of 257 female P. nianiculatus ob- of 41 females; and 16 to 31 July, 19.6 tained in the summer from the Black percent of 46 females. Hills and autopsied, 48.3 percent showed Adults molting from winter to sum- indications of reproductive activity. Be- mer pelage were taken as early as 10 cause many nearly-mature subadults car- June, but maximal time of pelage re- ried fetuses, both adults and subadults placement occurred from late June to were examined in this study. Of 48 fe- mid-July. Osgood (1909:19) indicated males collected in June, 16 were preg- that there is but one complete annual nant, having an average of 5.2 (4-7) change in pelage of adult Peromyscus embryos that were 9.3 (4-19) in crown- and that the process commences near the caudad. indi- rump length. Four placental scars were head, progressing Molting manifest in the reproductive tract of viduals trapped in late August and early another individual, and seven females September appeared to be acquiring the were lactating. The average length of characteristic long, dense pelage of win- testes of 67 males obtained in June was ter, and apparently were undergoing a

9.0 . In 24 of 87 females were second seasonal molt. The ( 6-18) July, high repro- in parturient, and carried 5.0 (4-9) fetuses; ductive rate of deer mice results many the average crown-rump length of the individuals of different ages occurring latter was 11.8 (2-25). Three other indi- simultaneously in the same area through- viduals had 4.3 (4-5) placental scars; a out warmer months, thus complicating like number of females had swollen uteri, interpretation of seasonal aspects of molt. possibly indicating recent implantation. Subadults undergoing post-juvenal molt Twenty additional females were lactat- were observed in all periods. The slate- ing. The average testicular length of 64 gray Juvenal pelage is replaced by males taken in July was 9.6 (6-18). Of brownish to buffy haii's that first occur 97 females obtained between 16 and 31 along the upper portion on each side. August, only 11 were gravid, containing Twenty-two kinds of ectoparasites 4.6 (4-6) embryos with an average were harbored by P. maniculatus from of 16.8 crown-rump length (6-27). the Black Hills: two lice, Hoplopleura Twelve other females had 4.3 (3-6) pla- hesperomijdis (Osborn) and Polyplax cental scars, and nine other individuals auricularis Kellogg and Ferris; three were lactating. Six females had swollen chiggers, Euschoengastia setosa (Ew- in uteri. Seventy-two males captured ing), Psuedoschoengastia farneri Lipov- the same period had an average testic- sky, and Xenacarus plnmosus (Green- In the first ular length of 9.8 (7-17). berg); four ticks, Dermacentor aiulersoni six of 25 females week of September, Stiles, Ixodes kingi Bishopp, /. spinipalpis contained 4.3 (3-6) young each that Hadwen and Nuttall, and individuals of had an average crown-rump length of the /. ochotonae-angustus complex; six 16.8 (8-23). Three females had 5.7 mites, Androhelaps fahrenholzi (Ber- Dermacarus En- (5-6) placental scars, and three other lese), hypudaei (Koch), laelaps stahidaris Laelaps mi- individuals were lactating. The aver- (Koch), croti (Ewing), Oniitlionysstis bacoti age testicular length of 27 males ob- (Hirst), and members of the genus Hirs- tained in the period 1 to 7 September tiomjssus; and seven fleas, Monopsyllus was 9.4 (5-18). Analysis of reproductive wagneri (Baker), M. eumolpi (Roths- data on a monthly basis masks short- child), Mahraeus telchinum (Roths- termed fluctuations. For the example, child), Orchopeas leucopus (Baker), O. of percentages pregnant individuals en- sexdentatus (Baker), CotaUagia decip- countered over four successive two-week iens (Rothschild), and Epitedia icen-

were: 1 to 15 Rothschild . periods June, 40.0 percent manni ( ) of 25 females; 16 to 30 June, 26.1 percent Comparison of 56 male and 42 female of 23 females; 1 to 15 July, 36.6 percent deer mice from the Black Hills revealed TURNER: MAMMALS OF THE BLACK HILLS 99 no significant secondary sexual dimor- blackish brown. When comparing habi- phism in external or cranial measure- tats of the l^adlands with those of the ments, although females are slightly Black Hills, Dice (1942:6) stated: "On larger in most dimensions. Males are the forested Black Hills, on the contrary, paler in tone (29.8±4.12) and greater in the variation from place to place in soil reflectance of reds than are color is relati\ and there is (50.6±0.36)+ cly slight, only females (27.4±4.85 and 49.5 0.28, re- a small amount of variability in the color spectively); however, there is consider- of deer-mice." In actuality, soils of the able variability in each of these param- Hills are exceedingly diverse in colora- eters. tion. For example, the two soils col- lected south of Peromijscus maniculaius is quite vari- Spearfish (Table 20) able in color and in external and cranial were within 20 yards of one another; dimensions in the Black Hills; this is sample A was obtained mid-way up a and B was taken at probably due to a greater divergence in steep slope, sample the base of the Soils from near topography, habitat, and edaphic factors slope. in the Hills than in the surrounding Rapid City register abnormally pale in tone due to a content that prairie. Dice (1941:16) noted a general high gypsum increases reflection of the soil. tendency for this species to increase in properties This in coloration of soils size westwardly over the Great Plains, great diversity invalidates correlations and indicated (1942:9) that the presence comprehensive of the Black Hills does not affect this between pelage and soil hues, unless a soil is taken at each site of trend in any significant manner. Deer sample cap- mice in the Hills are intermediate in size ture throughout the entire Hills region. soils of the Red between eastern, northern, and western Collectively, Valley "racetrack" differ from populations, but are assignable to the significantly those of the Limestone Plateau and Cen- western subspecies (Table 18). Pero- tral in tone and re- mijscus manicidatus nebrascensis differs Basin, being paler reds more Soils of the from P. m. hiteus to the east in being flecting intensely. Central Basin somewhat larger externally and cranially, and in average paler 13 and Table in tone than those of the Limestone Pla- being darker dorsally ( Fig. teau, but otherwise are simi- 19). Osgood (1908:78) noted that, al- they fairly lar. though typical nebrascensis average de- cidedly larger than typical liiteus, indi- Peromijscus manicuJatus in the Black vidual variation in each race is great Hills is much darker on the average than enough to allow overlap in the range of either the small "ochraceous-buff" race various parameters. For example, he re- on the arid plains south and east of the garded five of 13 specimens from Elk Hills, or the large "reddish" race on the the soils of eastern 13 Mountain as resembling Ititeus, and sandy Wyoming ( Fig. other eight as resembling nebrascensis. and Table 19). Collectively, specimens Occasional high frequencies of paler and from the southern Hills (especially from smaller mice in the foothills and periph- southwest of Minnekahta) are, on the due to eral areas may be expected gene average, paler than those from the north- with of the sur- exchange populations ern section, and specimens from the from Rapid rounding plains. Specimens western part (especially from the vicin- City, for example, approach the dimen- it>^ of Ditch Creek, Beaver Creek, Four sions of luteus rather than nebrascensis. Corners, and Buckhorn) have greater Stebler and Dice (1939:391-392) reflection of reds than individuals from (1942:1-10) both discussed the correla- the eastern Hills. tion between soil color and pelage color Measurements additional to those of P. maniculaius in the study area. In in Table 18 for 98 adult general, the soils of the Black Hills are given specimens the Black Hills are: dark in color, varying from reddish to from zygomatic 100 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

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Table 19.—Geographic variation in coloration of the mid-dorsal pelage of adult Peromysctts maniculatus obtained in summer from the Northern Great Plains.

Number and sex of specimens averaged TURNER: MAMMALS OF THE BLACK HILLS 103

(UMMZ); 16 mi NW Custer, 1 (UMMZ); cality, Saguache Valley, 20 mi W Sagauche, Harney Peak, 7240 ft, 4 (UMMZ); unspeci- Saguache Co., Colorado). fied 9 3 5 locality, (1 NRW, SDMT, UMMZ). Neotoma cinerea orolestes—Goldman, 1910, N. Custer Palmer 8 mi Hill County: Gulch, SE Amer. Fauna, 31:104, 19 October. Cit>', 5300 ft, 12 (FMNH); 3.5 mi S, 0.5 mi Neotoma grangeri J. A. Allen, 1894, Bull. Amer. W Ke\'stone, 5000 ft, 8; 5.75 mi N, 5.75 mi E Mus. Nat. Hist., 6:324, 7 November (type Custer, 5220 ft, 19; Roby Springs, 4 mi N, locality, Black Hills, Custer, Custer Co., 22 mi W Custer, 5400 ft, 2; Roby Canyon, 2 South Dakota). mi N, 22 mi W Custer, 5200 ft, 4; Squaw [Grace Coolidge] Creek, 14 (10 AMNH, 4 Bu-shy-tailed woodiats are boreal FMNH); Custer, 35 (32 AMNH, 3 FMNH); mammals of higher altitudes and lati- Harnev National mi Forest, 3-5 E Custer, 3 tudes of the northern and western United (UMMZ); 4 mi SW Custer, 9; Lightning States. In the Black Hills, Neotoma cin- Creek, 8 mi SW Custer, 1 (UMMZ); 4 mi NE erea crevices in Otis, 1 (UMMZ); 3 mi N Pringle, 5000 ft, 2 occupies rocky outcrops, (UMMZ); Campbell's Ranch, Elk Mountain, mine-shafts, and abandoned cabins from 4800 14 Wind Cave National ft, (USNM); the boreal cap down through the transi- Park, 10 (1 8 1 6 mi UW, UMMZ, WCNP); tion zone to about 4000 feet. These rats N, 1 mi E Wind Cave, Wind Cave National are active the around and are com- Park, 4400 ft, 1; 5 mi N, 2 mi E Wind Cave, year Wind Cax'e National Park, 4200 ft, 4; Buffalo mon throughout the study area. Vernon Corral, Wind Cave National Park, 4300 ft, 8; Bailey (1888:442) first noted this species 2.5 mi N, 5 mi E Wind Cave, Wind Cave in the Black Hills, indicating that its National Park, 3700 ft, 5; Beaver Creek Can- principal food seemed to be seeds of yon, Wind Ca\'e National Park, 4200 ft, 4; conifers and that of Wind Cave, Wind Cave National Park, 4; large piles gnawed Headquarters, Wind Ca\e National Park, 4100 cones often were scattered around en- ft, 8; Wind Cave Canyon, Wind Cave National trances to woodrat dens. Park, 4100 ft, 28; 11 unspecified locality, Woodrats are most abundant (UMMZ). Fall River Counti/: 1 mi N, 6 mi among and vertical crevices in E Hot Springs, 3400 ft, 2; 1 mi N, 5.5 mi E ledges rocky Hot Springs, 3400 ft, 2; 5.5 mi E Minnekahta, outcrops that form the steep sides of 4050 4 mi E Hot 3400 ft, 4; Springs, ft, 3; valley and canyon systems; examples are 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 72; Beaver Creek and Ditch Creek valleys, 1.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 6; and and Little Spearfish canyons. 1 mi S, 2 mi E Hot Springs, 3400 ft, 4; 2 mi Big Such areas of S, 2 mi E Hot Springs, 3600 ft, 2; 4 mi S Hot support growths ponderosa Springs, 1 (UMMZ); 3 mi N, 2 mi E Edge- pine and creeping juniper. At lower ele- mont, 3500 8 mi S Hot 3 ft, 1; Springs, vations, where rocky terrain is less com- (UXLMZ); Angostura Dam, 6. mon, buildings and other man-made fa- WYOMLNG: Crook County: 6.5 mi SSE cilities often are inhabited. Mine shafts Alva, Bear Lodge Mountains, 2 (UMMZ); and caves also are I have Devils Tower, flood plain Belle Fourche River, frequented; in at 33.50 ft, 2 (USNM); 15 mi ENE Sundance, observed N. cinerea Wind Cave 3825 Sand 1 3 mi ft, 11; Creek, (UW); NW depths of 325 feet below ground surface, Sundance, 5900 ft, 12; 2 mi NW Sundance, and 2300 feet from any known entrance 11; 1.5 mi NW Sundance, 5000 ft, 1; Sun- of the cave. dance, 10 (USNM). Weston Counttj: 1.5 mi E Buck-horn, 6150 ft, 2; 4 mi E Four Corners, This species sometimes is active in 2; 9 mi N, 1 mi E Newcastle, 5; 6 mi N New- late afternoon and one individual was 1 mi 4800 11 castle, 1; N Newcastle, ft, hrs taken by J. A. King at 16:30 (MST) (UMMZ); Newcastle, 12 (USNM). southeast of Hill on 29 March 1946. Additional records.—SOUTH DAKOTA: City This cricetid builds nests that are Pennington Counttj: Rochford (BB); Medicine large Mountain, 6200-6300 ft (Dice, 1942:3). WY- globular or cup-shaped masses composed Beaver OMING: Weston County: Stockade of dry grass, moss, leaves, and other fi- 4300 ft Creek, (Dice, 1942:3). brous materials; these nests generally contain cones of pine and spruce, bones Neotoma cinerea orolestes Merriam of various animals, and other debris. On Bushy-tailed Woodrat 4 July 1967, I found a mandible of Canis in a nest to the rear of Neotoma orolestes Merriam, 1894, Proc. Biol. lupus Davenport Soc. Washington, 9:128, 2 July (type lo- Cave, and a subadult in Wind Cave 104 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

amassed numerous flashbulbs and can- (Baker), and a chigger, Euschoengastia

its setosa . dles within nest. ( Ewing ) Black Hills woodrats were described Eight pregnant females taken in mid- A. Allen 1894a: as a June contained 3.6 (3-4) embryos that by J. ( 324-325) sepa- rate Neotoma based on had an axerage crown-rump length of species, grangeri, a series of 14 indi\iduals from Custer 32.1 (4-50). Three lactating females and two others from 11 of were captured in late July and two others Glendale; these were subadults. were collected in August. A female ob- specimens How- in his revision of the Neo- tained southwest of Sturgis on 14 August ever, genus tomcL Goldman indicated 1967 had six placental scars, and two (1910:105) others taken southwest of Lead in the that topotypes of N. grangeri did not diff^er from N. c. a con- same month had four placental scars each typical orolestes, clusion with which I concur. in the uterine walls. Testicular length of adult males varies seasonally. The a\'er- Some secondary sexual variation is age testicular length of four males taken apparent among the Black Hills popula- in June was 21 (16-30); whereas that of tions of N. cinerea. Females are paler in two captured in July was 12 each, and tone of color and greater in breadth of that of two obtained in August was 6 and interorbital constriction and length of are 7, respectively. Juveniles are common in maxillary tooth-row, but males collections made from mid-June through slightly larger in all other measurements. August. Two suckling young, \\hich Because individuals are quite ^•ariable, were only a few days old, were removed the sexes were not treated separately in from the teats of a female taken in Dav- subsequent analysis. enport Cave on 26 June 1967; the young Although no overall pattern of varia- were nearly naked and darkly pigmented tion in size is apparent, coloration of dorsally. Three suckling young ob- woodrats \'aries from place to place in tained on 8 July from northeast of Deer- the Hills. For example, specimens from field were older and in fine, short juvenal Beaver Creek are paler in tone (29.5± pelage that was dark gray in color. 6.26), reflect reds (51.9±0.27) more in- Change from j^nenal to subadult pelage tensely, and greens (23.7±0.13) and was evident in many specimens collected blues (24.2±0.20) less intensely than in late July, and several adults were un- do specimens from Ditch Creek (24.7 dergoing annual molt o\'er the sides and ±3.96, 45.2±0.27, 28.1±0.15, and 26.5 rump in late July and early August. ±0.29, respectively). Specimens from White, star-shaped spots were noted on other localities represent gradations be- the foreheads of many individuals, espe- tween these extremes. cially on subadults. Neotoma cinerea orolestes differs Chambers (1948:8) observed a red from N. c. rupicoh on the plains to the squirrel kill a woodrat in the northern south and cast in being darker and ha\- Hills near Lead. A female Neotoina ing somewhat larger external and cranial from west of Nemo and a male taken dimensions. Specimens from the Black Hills arc the southwest of Sturgis harbored ticks of on average significantly darker in tone and reflect blues more in- the Ixodes ochotonae-augustus complex. than do other of Indi\'iduals from southwest of Lead were tensely representati\es orolestes from west and nortli of the Hills parasitized externally by two ticks, (Fig. 14 and Table 21), but there were Ixodes spinipalpis Hadwen and Nuttall, no significant differences in external or and Dermacentor andersoni two Stiles, cranial measurements between these pop- mites, Andwhelaps fahrenhohi (Ber- ulations. Specimens from 0.5 mi E Buck- and amhulans lese) Haemogamasus horn. Weston Co., Wyoming, and a sub- two (Thorell), fleas, Orchopeas sexden- adult female from the Angostura Dam, tatiis (Baker) and Monopsylus ivagneri Fall River Co., South Dakota, are some- TURNER: MAMMALS OF TIIK BLACK HILLS 105

Little Spearfish Canyon, 2 mi S, H) mi W Lead, 0000 ft, 14; Little Spearfish Canyon, Savoy, 4 (USNM); Annie Creek, Black Hills, 6500 it, 1 (USNM); Deadwood, 2 (USNM); 2 mi W Nemo, 4700 ft, 2. Meade Counti/: Van- ocker Canyon, 3 mi S, 1 mi W Stiugis, 4600 ft, 1; Davenport Cave, 3 mi S, 0.5 mi W Sturgis, 4400 ft, 3. Pennington County: I3ark Canyon, 4 (AMNH); Diamond S Ranch, near Rapid City, 2 (UMMZ); Rapid City, 2 (1 NRW, 1 USNM); 15 mi SW Rapid City, 1 (NRW); 4.5 mi N, 2.5 mi E Deerfield, 4; Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 10; Ditch Creek, 14 mi W Hill Citv,

6400 ft. 10; 8 mi W Hill City, 7000 ft, 3 (UMMZ); Clendale, 2 (AMNH); Hill City, 1 (FMNH); Summit Peak, 3 mi SE Hill City, 5700 ft, 3 (UMMZ); Mushroom Rock, 4 mi SE Hill City, .5500 ft, 2 (UMMZ); Willow Creek, 4 mi SE Hill City, 5700 ft, 1 (UMMZ); unspecified locality, 1 (UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, 5300 ft, 1 (FMNH); Bull Springs, 2 mi N, 9 mi W Custer, 10 (UMMZ); Custer, 15 (2 USNxM, 12 AMNH, 1 FMNH); Housing Area, Wind Cave National Park, 4100 ft, 2; Wind Cave National Park, 7 (1 USNM, 1 WCNP, 5 UMMZ); Otis, 1 (UMMZ); Elk Mountain, 2 (USNM); unspecified locality, 7 (UMMZ). Fall River County: Angostura Dam, 1. WYOMING: Crook County: Devils Tower, 2 (USNM); Sand Creek, 1 (USNM); 15 mi N Fig. 14. variation in coloration of Geographic Sundance, 1. Weston County: 0.5 mi E Buck- the mid-dorsal pelage of three groups of sum- horn, 6100 ft, 5; 4 mi E Four Corners, 4; 3 mi mer-taken adult \\oodrats. Neotoma bushy-tailed N Newcastle, 1 (UW). cinerea orolestes averages much darker in tone than A^ c. rupicola, and specimens of N. c. orolestes from the Black Hills average much Clethrionomys gapperi brevicaudus darker than non-Black Hills representatives to (Merriam) the west and north. Red-backed Vole

Evotomys gapperi brevicaudus Merriam, 1891, what more pale than typieal Black Hills N. Amer. Fauna, 5:119, 30 July (type lo- 3 representatives, but are still best referred cality, mi N Custer, 6000 ft, Custer Co., South Dakota). to orolestes. Selected average external — and cranial measurements of eight males Clethrionomys gapperi brevicaudus Bole and Sci. Publ. Cleveland and 15 females from the Black Hills are: Moulthrop, 1942, Mus. Nat. Hist., 5:153, 11 September. total length, 359.1 ±20.67; tail length, 148.0± 10.31; hind foot length, 40.6 ± The red-backed \ole is distributed in 2.12; ear length, 33.4±2.55; weight, 272.5 boreal and montane coniferous forest and ±77.95; greatest length of skull, 48.3 ± its successional stages, and in brushy ri- 1.82; zygomatic breadth, 25.8±1.06; mas- parian habitats of North America. It toid breadth, 20.2 ±0.63; rostral length, inhabits the boreal cap of the Black 20.1 ±0.84; interorbital constriction, 5.9 Hills down to eknations of 4600 feet, ±0.28; length of incisive foramen, 11.2± but is most numerous above 5200 feet. 0.63; length of maxillary tooth-row, 9.9 It is active the year around. Although ±0.49; depth of skull, 16.5 ±0.61. common in the Hills, no specimens are at hand from Meade or Fall Ri\'cr coun- Specimens examined (L34).—SOUTH DA- ties. KOTA: Lawrence County: Little Spearfish Canyon, 2 mi S, L3 mi W Lead, 6000 ft, 1; Merriam named the subspecies, 1)revi- 106 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Table 2L—Geographic variation in coloration of the mid-dorsal pelage of adult Neotoma cinerea ol:)tained in summer from the Northern Great Plains.

o O o Number and sex c B C a; O o a> 0; of specimens o OJ ,s

23 (85,15$) TURNER: MAMMALS OF THE BLACK HILLS 107

duced annually. A subadult female taken tinct. Adults taken in mid-June and early along Ditch Creek on 10 June 1965 car- July frequently were molting into the ried three embr\os that had a crown- short grayish- to reddish-brown summer rump length of 4 each. Two adult fe- pelage; new hairs were enshrouded be- males obtained about this time showed neath the overlying older pelage on both no signs of reproductive actixity. The av- sides and dorsum. Because a distinctive erage length of testes of nine males ob- moltline is not formed the process of tained in June was 9.0 (7-10); corre- pelage replacement is rather impercepti- sponding measurements of five subadults ble. Fur of juveniles is more grayish in

were 7.8 (6-9) . Eight of 15 adult females color than that of adults. Many sub- captured in July were pregnant; they had adults taken in summer were molting an average of 6.8 (5-9) embryos that simultaneously over much of the body, were 10.1 (5-20) in crown-rump length. whereas others had completed post- Testicular length of 12 males collected juvenal molt and were in fresh brownish in July was 9.7 (7-11); that of three sub- pelage. adults was 8.3 (7-10). A lactating adult A male captured northeast of Custer female was captured southwest of Rock- on 19 June 1967 was infested with ticks, erville on 19 August 1965 and testicular Dermacentor anclersoni Stiles. length of six adult males obtained in Comparison of 27 male and 22 fe- August was 6.8 (6-8). Strangely, observa- male C. g. brevicaudus revealed no ap- tions concerning the occurrence of pla- parent secondary sexual dimorphism in cental scars in C g. hrevicaudus are coloration or in external and cranial di- lacking in field notes of collectors, even mensions, nor was a pattern of variation though notations of this type are entered in these characters evident in the Black for all other species; perhaps placental Hills. Clethrionomys gapperi hrevicau- scars are more ephemeral in red-backed dus differs significantly from C. g. galei voles than in other small mammals. to the west in Wyoming in having a Winter pelage is longer and softer shorter tail and ear, longer hind feet, than that of summer and the reddish a higher skull, and darker pelage that brown dorsal stripe is much more dis- reflects reds more intensely (Table 22).

Table 22.—Geographic variation in coloration of the mid-dorsal pelage of adult Clethrionomys gapperi obtained in summer from the Northern Great Plains.

Number and sex of specimens averaged 108 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

3 mi SE Hill 5300-.5400 11 Clethrionomys gapperi hremcaudus also Gulch, City, ft, (UMMZ); Willow Creek, Nelson's Place, 4 mi averages somewhat shorter in total SE Hill City, 5300-5400 ft, 13 (UMMZ); and in lengths of skull length greater Harney Peak, 5 mi SE Hill City, 7240 ft, 1 in lamb- and rostrum, and zygomatic and (UMMZ); Rockerville, 1 (UMMZ); 3 mi S, doidal breadths (Table 23). From C. g. 1 mi W Rockerville, 3; Keystone, 1 (NRW); 17 mi NW Custer, 11 (UMMZ); 16 mi NW loringi to the north and east, brevicaiiclus Custer, 22 (UNLMZ); 16 mi SW Rapid Citv, 1 differs significantly in longer hind feet, (UMMZ); unspecified locality, 2 (UMNIZ). lambdoidal breadth and a greater higher Custer County: Palmer Gulch, i mi SE Hill skull; it is also smaller in total length City, 5300 ft, 9 (FMNH); 0.5 mi E Sylvan 62.50 1 3.5 mi 0.5 mi and length of tail, and greater in lengths Lake, ft, (UMMZ); S, W Keystone, 5000 ft, 1; 5.75 mi N, 5.75 mi E of ear, skull, and rostrum and in zygo- Custer, 5220 ft, 21; 3 mi N Custer, 6000 ft, matic breadth. Measurements additional 2 (USNM); Custer, 21 (18 AMNH, 3 FMNH); to those given in table 23 for 49 adult Bull Springs, 2 mi N, 9 mi W Custer, 7 red-backed voles from the Black Hills (UMMZ). are: weight, 25.4±;4.48; interorbital con- WYOMING: Crook County: 3 mi NW 5900 Rattlesnake 3 striction, 3.9 ±0.13; length of incisive Sundance, ft, 3; Creek, (USNM). Weston County: 1.5 mi E Buck- foramen, 4.9 ±0.26; length of maxillary horn, 6100 ft, 21; 12 mi SE Newcastle, 1 tooth-row, 5.4 ±0.25. (UMMZ). Specimens examined (229).—SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- Microtus longicaudus longicaudus 6 mi S, 2 mi W 4600 ft, 6; yon, Spearfish, (Merriam) Tinton, 5900 ft, 1; 2 mi S Tinton, 6100 ft, 22; Deadwood, 1 (USNM); Dumont, 6100 ft, 5 Long -TAILED Vole (USNM); 2 mi W Nemo, 4700 ft, 4; Nemo, Arvicola (Mijnomes) longicaudus Merriam, 4700 ft, 1. Pennington County: Jim Creek, 1888, Amer. Nat., 22:934, October (type east of Merritt, 1 (NRW); Rapid" Creek, 1.5 locality, Custer, 5500 ft, Custer Co., South mi W Rochford, 1 (UMMZ); Diamond S Dakota . Ranch, near Rapid City, 1 (UMMZ); Beaver ) Microtus — A. Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, (Mynomes) longicaudus T- Allen, Bull. Amer. Mus. Nat. 21; 3 mi N, 7 mi W Deerfield, 6900 ft, 1; 1895, Hist., 7:266, 21 1 mi S, 2 mi E Deerfield, 6400 ft, 1; McVey August. Burn, 1 (NRW); 20 mi N Elk Mountain, 6000 The long-tailed \'ole occurs on the ft, 2 (USNM); Ditch Creek, 14 mi W Hill boreal of the Black Hills and inhab- City, 6400 ft, 3; Hill Citv, 1 (AMNH); Palmer cap

Table 2.3. —Geographic variation in selected external and cranial measurements of adult Clefliriono- mys gapperi from the Northern Great Plains.

Number and sex of specimens TURNER: MAMMALS OF THE BLACK HILLS 109 its riparian environs bordering cold in association with long-tailed voles are streams that descend into the lower semi- Peroniyscus nianiculatus, Eutamias mini- arid transition zone. It is widely dis- mus, Zapus hudsonius, and Sorex cine- tributed but none has been taken as yet reus. in Fall Ri\er County. Although speci- Reproducti\e data for long-tailed mens have been obtained in the Hills voles in the Black Hills are adecjuate only as low as 4200 M. is feet, longicaudus for July. Of two females taken west of most abundant at elevations above 6000 Hill City in mid-June 1965, one carried feet. The Black Hills is population iso- five embryos which were 10 in crown- lated from other montane generally pop- rump length; the other showed no signs ulations westward and southward in Wy- of reproductive activity. Testicular The does not hibernate oming. species length of a male obtained on 28 June and indi\'iduals have been in captured 1967 was 12. Fifteen of 23 females cap- all seasons. tured in July were pregnant; there was an A distinct pattern of ecological sepa- average of 4.7 (4-6) embryos in each, ration between M. longicaudus and M. which were 10.3 (3-24) in crown-rump pennsyJvanicus is not readily discernible length. One female collected northwest in the Hills. W. W. Granger wrote about of Deerfield on 8 July 1965 had eight them as follows, "I found these mice placental scars; another taken east of [A/. pen}isyJvanicus] in the same locali- Buckhorn on 18 July 1951 was lactating. ties as the other species [M. longicau- An additional female trapped southwest dus]. Some were caught on a hillside of Lead on 30 July 1968 had a swollen which was covered with aspens, and the uterus, possibly indicating recent im- rest along the banks of a creek" (J. A. plantation. Testes of 12 adult males cap- Allen, 1895a: 267). Both species fre- tured in July had an average length of quently are captured in the same trap- 11.2 (10-12). On 9 August 1967, two line in the mesic Northern Hills. For females were captured south of Spear- example, a Sherman live-trap placed be- fish; one carried six embryos that were neath the overhanging bank of Little 4 in crown-rump length, whereas the Spearfish Creek in the Timon Camp- other was lactating and had seven pla- ground obtained one M. longicaudus and cental scars. Juvenile and subadult long- one M. pennsylvanicus within an hour tailed voles are common in collections 19:30-20:30 hrs on 31 1968. made and and ( MDT) July throughout July August, Vegetation in the vicinity of the trap many of the pregnant females were sub- was composed of mosses, grasses, and adults. deciduous shrubs. Five young voles taken in Beaver At higher elevations, M. longicaudus Creek Valley in early July were in Juv- ranges farther from riparian situations enal pelage, which was fine, soft, and than does M. pennshjvanicus, which in- dark. Twelve juveniles collected through- habits moist meadows and edges of out July either had completed post- spruce and aspen forest. In the latter juvenal molt, or replacement of juvenal habitat, long-tailed \'oles overlap Cleth- pelage was in process with new hairs lionomys gapperi. At lower elevations, exposed beneath the old pelage on the M. longicaudus is more restricted to sides and dorsum. In the same period, streamside environments and lowland three subadults were molting to an adult marshes of rushes, sedges, and cattails, pelage. In adults, change from old, or other moist areas of much brush and faded winter pelage to fresh summer fallen logs. In these areas, M. pennsyl- pelage was noted in late June and early vanicus ranges farther from riparian situ- July; irregular patches of new hair oc- ations and is more tolerant of the drier curred on the rump and over the shoul- meadows. In addition to M. pennsylvani- ders. cus and C. gapperi, other species trapped A male trapped in Beaver Creek 110 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

ments to in Table Valley on 8 July 1965 harbored ticks of additional those given the Ixodes ochotonae-angustus complex. 24 for 31 adults from the Black Hills are: red Significant secondary sexual dimor- percent reflectance, 50.0±0.23; phism was not evident in comparison of percent green reflectance, 26.7 ±0.18; 14 adult males with 17 adult females percent blue reflectance, 23.3±0.16; from the Black Hills, although males weight, 41.9±7.04; interorbital constric- of incisive fora- are slightly larger in most measurements tion, 3.8±0.15; length and females (50.9 ±0.25) reflect reds men, 5.0±0.23. more intensely than males (48.8±0.15). Specimens examined (172).—SOUTH DA- A distinct pattern of variation in colora- KOTA: Lawrence County: Big Spearfish Can- 3 mi S 4200 5900 tion or size within the Hills populations yon, Spearfish, ft, 2; Tinton, ft, 1; 2 mi S Tinton, 6100 ft, 4; Little Spear- is not discernible. fish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, noted that M. /. Bailey (1900:48) 3; Cheyenne Crossing, 8 mi SW Lead, 1 longicaudus and M. I. mordax are "very (SDSU); Boxelder Creek, Steamboat Rock similar" in external and cranial dimen- Campgrounds 1 (NRW); Dumont, 1 (USNM). Meade 7 mi S, 1 mi W Sturgis, 4700 sions. After examining specimens from County: 1. 1.5 Idaho and Long ft, Pennington County: Rapid Creek, throughout Wyoming, mi W Rochford, 4 (UMMZ); Beaver Creek, mordax in the (1965:655) placed synon- 4 mi N, 10.5 mi W Deerfield, 6400 ft, 17; that omy of longicaudus, pointing out Ditch Creek, 14 iTii W Hill City, 6400 ft, 2; 1 20 mi Elk C. H. Merriam selected an atypically Redfern, (USNM); N Mountain, 6000 ft, 12; 17 mi NW Custer, 7 (UMMZ); dark vole with long ears as the holotype. 16 mi NW Custer, 23 (UMMZ); Palmer Gulch, When 31 voles from the Hills long-tailed 3 mi SE Hill City, 5300-5400 ft, 7 (UMMZ); were compared with 29 from north- Nelson's Place, 4 mi SE Hill City, 5300-5400 2 5 mi Hill central Wyoming (Big Horn, Fremont, ft, (UMMZ); Palmer Gulch, SE 2 5 Johnson, Sheridan, and Washakie coun- City, (UMMZ); unspecified locality, (UMMZ). Custer County: Palmer Gulch, 8 ties southeastern ) and 80 from Wyoming mi SE Hill City, 5300 ft, 4 (FMNH); 16 mi (Albany, Carbon, Converse, Laramie, W Custer, 7 (USNM); Custer, 6 (2 USNM, 4 and Natrona counties); the former aver- AMNH); 4 mi E Custer, Harney National 1 Elk 4900 aged darker in tone and possessed longer Forest, (UMMZ); Mountain, ft, 4 (USNM); 4 mi SW Custer, 1; 18 mi SW ears and maxillary tooth-rows (Table Custer, 5300 ft, 5 (USNM); Bull Springs, 2 24). mi N, 9 mi W Custer, 8 (UMMZ). Even though the pelage of M. longi- WYOMING: Crook County: Bear Lodge 6000 1 caudus is extremely uniform in reflect- Mountains, Warren Peak, ft, (USNM); Bear Lodge Mountains, 6.5 mi SSE Alva, 1 ance of various hues, this species is quite (UMMZ); 3 mi NW Sundance, 5900 ft, 7; variable throughout its range in all other 1.5 mi NW Sundance, 5000 ft, 1; Sundance, For relative to the parameters. example, 6000 ft, 12 (USNM); Rattlesnake Creek, 6000 Black Hills population, 28 individuals ft, 2 (USNM). Weston County: 1.5 mi E 6150 4 mi E Four 1. from Carbon County, Wyoming, are Buckhorn, ft, 15; Corners, Additional record.—SOUTH DAKOTA: darker in tone of color (21.0±1.98), 17 Pennington County: Rochford (BB). specimens from Natrona County have a similar of ear 15.8± average length ( Microtus ochrogaster haydenii (Baird) 1.59), and single representatives from Prairie Washakie and Sheridan counties each Vole have longer maxillary tooth-rows (7.5 Arvicola (Pedomys) haydenii Baird, 1858, in

of and surveys . . . and 7.0, respectively). Thus, the above- Reports explorations from the Mississippi River to the Pacific mentioned apparent differences attribut- . . lo- Ocean ., 8(1):543, 14 July (type able to the Black Hills of M. population cality, Fort Pierre, Stanley Co., South Da- longicaudus easily fall within the range kota). of variation of populations southward Microtus ochrogaster haydenii—Osgood, 1907, Biol. 18 and westward of the Hills, and I must Proc. Soc. Washington, 20:48, April. concur with the findings of Long {loc. cit. . ) Average color and cranial measure- Microtus ochrogaster, in contrast to TURNER: MAMMALS OF THE BLACK HILLS 111

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H<; 112 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY other members of the genus that occur rushes, and sedges; Microtus longicau- were ob- in the study area, is relatively tolerant dus and M. pennsylvanicus of xeric conditions and is widely distrib- tained in these areas, but Cleithriono- uted wherever suitable grassy habitats mys no longer was taken. Below 4000 are available on the Great Plains of cen- feet, upward extensions of the xeric tral North America. Prairie \'oles have prairie were encountered as e\ idenced been taken in all counties within the by the presence of thistles, soapweed, Black Hills, and are found in the drier mustards, and an occasional pricklypear Prairie voles were taken under valley systems that penetrate the foothill cactus. transition zone. They occur up to 5000 these conditions, whereas the other three encountered. feet, but are most abundant below 4400 microtines were no longer feet along the interface of the surround- Fifty-eight individuals were captured southwest of Minnekahta 13-18 ing prairie. Wasteland, dry ditches along ( June that form the southern roadways, and lightly grazed pastures 1967) on slopes also are inhabited by this vole. Vernon limits of the Black Hills. The area con- Bailey (1888:445) reported the first sists of arid grassy pastureland, expanses specimens of M. ochrogaster in the Black of dryland farming (mostly small of Hills region. grains), and a series ridges upon which and western red From 7 to 11 August 1967, a survey ponderosa pine cedar Most of the voles were was taken along the floor of Big Spearfish grow. taken fencerows agri- Canyon, out onto the adjacent prairie. along bordering cultural in of The vegetation of this moist northern fields, vegetation composed brome cord west- canyon system is rather diverse. At bluegrass, grass, grass, ern wheat and scattered forbs. higher elevations, ponderosa pine is grass, Vole were well-defined but lo- fairly restricted to the arid bluffs and runways and of were white spruce dominates the mesic can- calized, piles grass cuttings common. collected near Hot yon bottoms, along with rushes, sedges, Specimens were on and in ap- dense stands of tall grasses, deciduous Springs grass ridges orchards that flank the Fall River. shrubs, and quaking aspen; Clethriono- ple extends rnys gapped, Microtus longicauclus, and Wind Cave National Park M. pennsijlvanicus were taken in these through the gently rolling foothills of the areas. Descending along the canyon, transition zone, grading into the more conditions gradually became less mesic. rugged topograph)' of the southeastern Deciduous elements such as bur oak, portion of the Black Hills. Here, where willow, paper birch, quaking aspen, box- Al. ochrogaster is one of the commonest elder, bunchberry, and alder are abun- small mammals, the mixed-grass prairie dant. The understory of parklike wood- of the plains intermingles with the pon- lands and small meadows is composed derosa pine of the higher elevations. of wild wild ameri- gooseberry (Rihes missouriense) , Chokecherry, plum (Prunus wild strawberry (Fragaria virginiana), cana), wild rose, hackberry, and sage- with chokecherry, poison ivy, Solomon's seal, brush are interspersed prairie starry solomonplume {Smilacina stel- grasses and forbs. Other mammals lata), yellow lady's slipper (Cijpripe- trapped in the runways of this species (Uiim sp. ), white geranium (Geranium were Pcromyscus maniculatus, Perogna- bicknellii), black-eyed susan, sunflower, thus hispidus, Mus musculus, and Reith- burdock (Arctium minus), cinquefoil rodontomys megalotis. and (Potentilla sp.), meadow sweet Reproductive activity was substantial

lucida . ( Spiraea ) Along Spearfish Creek, from late spring throughout the summer, and adjacent moist areas at these inter- and continued at least into early autumn. mediate elevations are snowberry, horse- Three females captured in late March tail (Equisetum sp.) flowering fern (Os- and early April carried 4.3 (4-5) em- munda sp.), wood fern (Dryopteris sp.). bryos that had an average crown-rump TURNER: MAMMALS OF THE BLACK HILLS 113

c> in adulls s(^ems to length of 5.7 (2-10). Reproducti\ data replacement be more are unavailable for May, early June, and erratic than in subadults and possibly of time. July (except for one female captured on extends over a greater period 26 July 1968 that carried three embryos Prairie voles from the Hills are heav- which were 8 in crown-rump length). ily infested \\ilh various types of ecto- Field work dining these periods \\'as un- parasites as follow: a louse, Iloplopleiira dertaken at higher ele\ations where acanthopus (Burmeister); a tick, Ixodes prairie voles do not occur, thus explain- spinipaipis Hadwen and Nuttall; two ing the lack of data. Sixteen of 26 fe- fleas, Orchopeas leucopus (Baker), and males taken in the last half of June were MonopsyUus ica^neri (Baker); two chig- pregnant, containing 3.5 (1-6) embryos gers, Euiromhicxda alfreddugesi (Oude- that had an a\ erage crown-rump length mans), and Euschoengastia setosa (Ew- of 10.5 (3-20). Two other females had ing); and seven mites, Dermacams hy- 3.0 (2-4) placental scars, and another pudaei (Koch), Mycoptes musculinus female had an enlarged uterus, which (Koch), Haemolaelaps jahrenhohi Ber- possibly indicates recent implantation. lese, Androlaelaps jahrenhohi (Berlese), A\erage testicular length of 22 adult Laelaps microti (Ewing), Ornithonyssus males obtained in this period was 11.0 hacoti (Hirst), and Haemogamasus am- (7-17). In the latter part of August, hulans (Thorell). three of nine females examined contained Examination of 26 adult male and 23 number of fetuses fetuses. The average adult female M. ochrogaster from the was 3.7 (3-4); the average crown-rump Black Hills revealed no secondary sexual 11.3 length of the fetuses was (6-20). differences in coloration, external meas- Two other females had five placental urements or most cranial dimensions. each. The of the scars average length However, females (6.9±0.32) are sig- testes of 11 males obtained in was August nificantly greater than males (6.6d=0.29) Five of females 13.0 (10-18). eight cap- in length of maxillary tooth-row. A pat- tured in early September were pregnant, tern of variation within the Hills was not and contained an average of 3.8 (3-5) evident. When comparisons were made that were 15.2 in crown- embryos (8-22) with populations from other areas in scars were rump length; four placental South Dakota and Wyoming, the great in the tract of one manifest reproductive variabifity of M. ochrogaster became ap- testicular other female. The average parent (Tables 25 and 26). Characters males obtained about length of five adult seem to vary independently and no def- and this" time was 12.6 (9-17). Juveniles inite trends are discernible, although in collections subadults are common specimens from the Hills tend to be made from April through September; darker in tone of color and relatively the females were several of gravid nearly large in external and cranial dimensions. mature subadults. Measurements additional to those given Due to the high reproductive rate of in Table 26 for 44 adult prairie voles molt were are: hind foot this species, seasonal aspects of from the Hills length, difficult to discern. Juveniles were in a 20.2±0.94; lambdoidal breadth, 12.4db fine dark pelage, although many were 0.39; interorbital constriction, 4.0 ±0.13; commencing the post-juvenal molt. As length of incisive foramen, 5.0+0.32; and voles attained about three-quarters of length of maxillary tooth-row, 6.7 it 0.34. adult size and adult pelage replaced Specimens examined (214).—SOUTH DA- be that of the subadults, new hairs could KOTA: Lawrence County: Big Spearfish Can- the 3800 6. observed underlying the older fur on yon, 1.5 mi S, 0.5 mi E Spearfish, ft, Meade Black Hawk, 22 (AMNH). dorsum and sides, and occasionally on County: Pennington County: 8 mi NE Rapid City, 4 the venter. Irregular patches of molt (UMMZ); 4 mi N Rapid City, 2 (AMNH); from were noted on adults in all months South Canyon, 3 mi W Rapid City, 13 (6 7 Rapid City, 9 (4 USNM, D which specimens were examined. Pelage AMNH, NRW); 114 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Table 25.—Geograpliic variation in coloration of the mid-dorsal pelage of adult Microtus ochrogaster obtained in summer from the Northern Great Plains. TURNER: MAMMALS OF THE BLACK HILLS 115

Table 26.—Geographic variation in selected external and cranial measurements of adult Microtiis ochrogaster from the Northern Great Plains.

Nimiber and sex of specimens averaged 116 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

southwest of on 25 of that species). Morris (1969:299) re- Rapid City August 1952 contained in ported an almost complete habitat sepa- eight (21 crown-rump ration between M. penmijlvanicus and length). A subadult female taken in Park on 1 Clethrionomys gapperi in Saskatchewan, Wind Cave National Septem- 1968 three with the former being confined to grass- ber was gravid, carrying land and the latter to aspen woodland. embryos that were 27 in cro\\'n-rump of the other above-men- In the Hills, however, both species fre- length. Many tioned females were ma- quently were taken in the same habitat. pregnant nearly are common Although M. pennsylvanicus was not ture subadults. Juveniles in collections made from captured in stands of aspen or other June through in other woodlands, C. gapperi often was ob- August, and undoubtedly peri- ods as well. tained in grassland and riparian situa- tions where brush, woodpiles, and fallen The process of pelage replacement logs were common; here, the two species in M. pennsylvaniciis is similar to that occurred side by side. At lower eleva- described for other microtines in the tions, where the meadow vole and prairie Black Hills. Adults molt from pale, vole occur together. M. penmijlvanicus faded winter pelage to fresh, dark sum- inhabits the lower, wetter areas and M. mer pelage in late June and July. All ochrogaster the drier places. For ex- stages of molt, from commencement to ample, in Wind Cave National Park, the completion, can be observed in adults meadow vole occupies the immediate taken on the same day at the same lo- brushy banks of Beaver Creek, whereas cality in mid-summer. Juveniles were in the prairie vole occupies the gi-assy up- a fine, dark fur that was replaced by land adjacent to the creek. In addition subadult pelage. Developmental pelage to the above named species, Peromijscus replacement depends on chronology and maniculatiis, Eiitamias minimus, Zaptis rate of maturation of the individual; thus, litidsoniiis, and Sorex cinereus were taken concurrent existence of several different- in the same ti'aplines with M. pennsyl- aged litters tends to complicate interpre- vanicus. tation of precise seasonal aspects of molt. In the Black M. Reproductive data for meadow voles Hills, pennsylvaniciis Der- in the Black Hills are available only for are parasitized externally by ticks, macentor andersoni Ixodes the period from mid-June through early Stiles, spini- Hadwen and and mem- August. Bailey (1924:528) indicated palpis Nuttall, bers of the Ixodes that parturient female M. pennsylvaniciis ochotonae-angustae mi- have been taken in all seasons. Four of complex, by laelaptid mites, Loehps and six females captured in late June carried croti (Ewing) Androlaelaps fahren- holzi and a En- 5.8 (5-7) embryos that had an average (Berlese), by chigger, setosa crown-rump length of 17.3 (7-27). One schoengastia (Ewing). scars. first other individual had six placental J. A. Allen (1877:176) reported The average testicular length of 17 adult meadow \'oles from the Black Hills area in 12.8 Arvicola and males obtained June was ( 10-15). under the name riparius, Thirty-four of 50 females captured in later (1894b: 347) assigned Hills speci- July were pregnant; the average number mens to a new subspecies, insperatus. of 5.9 2-9 and their aver- his revision of the embiyos was ( ) , In genus Microtus, age length was 12.1 (4-27). Four other V. Bailey (1900:20) listed Arvicola in- females were lactating and another four speratus in the synonomy of M. p. mo- had enlarged uteri, possibly denoting destus, but later (1920:72) included recent implantation. The average length Black Hills specimens under the name of testes of 23 males obtained in July M. p. icahema described from eastern was 14.4 (10-18). A female captured Montana and western North Dakota: north of Sundance on 15 August 1951 R. M. Anderson (1943:92) returned in- carried six fetuses and another obtained speratus to subspecific rank, concluding TURNER: MAMMALS Ol^^ THE BLACK HILLS 117 that icahenia was not distinct. S. Ander- Microtus pennsylvanicus insperatus is son (1956) discussed subspeciation in evid(>ntly quite variable throughout its the meadow vole in the western part of geographic range in external and cranial its range. dimensions, but each character seems to Examination of 31 adult male and 37 \ary independently and no trends are ap- adult f(^male M. pennsylvanicu.s from parent (Table 27). Meadow voles from the Black Hills in the Black Hills revealed no secondary are much darker com- sexual variation in coloration. Males are parison to specimens from Wyoming and but voles from significantly larger than females in lamb- Montana, Harding doidal breadth (12.3±0.46, 11.9±0.37) County, South Dakota, approach the and length of incisixe foramen (5.5± color of those of the Hills in tone. Meas- urements additional to those in Table 0.31, 5.3±0.20), and are shghtly larger 27 for 68 adult M. in most other dimensions. A pattern of pennsijlvanicus from variation in external and cranial dimen- the Black Hills are: percent red reflect- sions is not discernible within the Hills ance, 51. 2 ±0.24; percent green reflect- but four specimens from west of Nemo ance, 25.9 ±0.20; percent blue reflect- 22.6 hind 20.8 (15.0 ±1.78) and six from northeast of ance, ±0.24; foot length, interorbital Custer (15.1 ±1.28) are darker in tone ±3.83; weight, 48.6±7.44; of color on the average than 14 individ- constriction, 3.6 ±0.15; length of incisive of maxil- uals from east of Buckhorn (18.6±:1.42), foramen, 5.4±0.27; and length five from northeast of Sundance (18.3±: lary tooth-row, 6.7±0.21. 27 from south of Tinton (17.3± 2.17), . — Specimens examined ( 301 ) SOUTH DA- six 1.87) and from Beaver Creek Valley KOTA: Lawrence County: Little Spearfish

(16.5±1.18). Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 4;

Table 27.—Geographic variation in selected color, and external and cranial measurements of adult Microtus pennsylvanicus from the Northern Great Plains.

Number and sex of specimens averaged 118 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

2 2 mi S Tinton, 6100 ft, 52; Roubiax Lake, systems of the Northern Great Plains and 2 mi 4700 ft, 6. Penning- (NRW); W Nemo, is common on nearly all streams that ton Countt/: Rapid City, 1 (NRW); Rapid originate in the Black Hills. Although Creek, 1.5 mi W Rochford, 3 (UMMZ); 2 mi few of this micro- W Oreville, 5500 ft, 1 (UMMZ); Beaver Creek, specimens semiaquatic 4 mi N, 10.5 mi W Deerfield, 6400 ft. 18; tine are available, I have observed musk- Beaver Creek, 3 mi N, 9 mi W Deerfield, 6400 rats throughout the Hills. Dens and sub- ft, 6; Castle Creek, 6500 ft, 7 (UMMZ); Bald merged trailways interlace the shorelines Hills, Pactola, 3 (USNM); 6 mi S, 9.5 mi W of lakes and reservoirs, and Rapid City, 3; Sheridan Lake, 2 (NRW); penetrate Spring Creek, south Rapid City, 1 (MHM); banks of all water courses of the region. 4 mi NW Hill City, 2 (UMMZ); 20 mi N Elk In the past, Ondatra undoubtedly has 6000 5 22 mi Mountain, ft, (USNM); Moon, been one of the most commercially im- W Hill City, 6200 ft, 7; Ditch Creek, 14 mi portant fur-bearers sought by trappers W Hill City, 6400 ft, 1; Hill City, 1 (AMNH); in the Black 1 Hills, but more recently, near Palmer Culch, ( UMMZ ) ; Palmer Gulch, 3 mi SE Hill City, 5300-5400 ft, 25 (UMMZ); muskrat pelts ha\'e fetched relatively 4 mi SE Hill 5300-5400 Nelson's Place, Citv, poor prices. Burrows excavated in dams ft, 10 (UMMZ); 16 mi NW Custer, 3 and irrigation ditches cause extensive (UMMZ); Horse Thief Lake, near Mount to these structures and lend a Rushmore, 1 (NRW); imspecified locality, 3 damage (UMMZ). Custer County: Palmer Gulch, 8 negatixe dimension to the economic im- mi SE Hill City, 5300 ft, 14 (FMNH); 3.5 mi portance of this species. 0.5 mi 5000 0.5 mi E S, W Kevstone, ft, 12; Compared to O. z. osoijoosensis to the Sylvan Lake, 6250 ft, 1 (UMMZ); 5.75 mi N west in Wyoming, O. z. cinnamominus is 5.75 mi E Custer, 5220 ft, 13; Custer, 5 smaller with a shorter (AMNH); Harney National Forest, 4 mi E cranially, notably is in Custer, 2 (UMMZ); 3 mi N Pringle, 5000 ft, maxillary tooth-row, and paler color, 3 mi (UMMZ); Lightning Creek, 8 SW Custer, being more reddish or buflPy instead of 5100 ft, 1 (UMMZ); Beaver Creek Canyon, brownish (Long, 1965:662). The species Wind Cave National Park, 4200 ft, 8; Head- was first reported from the Black Hills quarters, Wind Cave National Park, 4100 ft, 1; Wind Cave Canyon, Wind Cave National by Bailey (1888:445). Park, 4100 ft, 1; Bull Springs, 2 mi N, 9 mi W Reproducti\'e and natural history data 9 Custer, (UMMZ). are lacking for this microtine in western WYOMING: Crook Bear County: Lodge South Dakota, but in northern Nebraska, Mountains, 2 (USNM); Bear Lodge Mountains, the muskrat breeds from late to 6.5 mi SSE Alva, 2 (UMMZ); 15 mi N Sun- — April dance, 5500 ft, 3; 15 mi ENE Sundance, 3825 early September adult females produce 3 ft, 6; mi NW Sundance, 5900 ft, 1; 1.5 mi one to four litters per year, averaging NW Sundance, 5000 ft, 4; Sundance, 15 about six voung per litter (Jones, 1964: (USNM); Ratdesnake Creek, 6000 ft, 1 235). (USNM); Sand Creek, 3750 ft, 2 (USNM). Weston County: 1.5 mi E Buckhorn, 6150 ft, Specimens examined (24).—SOUTH DA- 25; Newcastle, 2 (USNM). KOTA: Lawrence County: Little Spearfish Additional recorrf.s.—SOUTH DAKOTA: Canyon, Savoy, 4 (USNM). Pennington Lawrence County: Spearfish (BB). Penning- County: Rapid City, 2 (1 USNM); Diamond ton County: (BB). Custer County: French S Ranch, near Rapid City, 2 (UMMZ); Spring Creek, 3 mi E Custer (Stebler, 1939:390). Creek, south of Rapid City, 2 (MHM); Tiger- ville, near Hill Citv, 1 (USNM); Hill Cit\-, 3 Ondatra zibethicus cinnamominus (AMNH); Harney National Forest, 1 (UMMZ). (Hollister) Custer County: Custer, 8 (7 AMNH, 1 FMNH). Fall River County: 1 mi N Hot MUSKRAT Springs, 1 (UW). Additional records.—SOVTH DAKOTA: Fiber zibethicus cinnamominus Hollister, 1910, Custer County: French Creek, 3 mi E Custer Proc. Biol. Soc. Washington, 23:125, 2 Sep- (Stebler, 1939:390). Fall River County: Hot tember (type locality, Wakeeney, Trego Co., Kansas). Springs (Woodward, 1930:109). Ondatra zibethicus cinnamomina— Miller, 1912, Family MURIDAE—Old World Bull. U. S. Nat. Mus., 79:232, 31 Decem- ber. Rats and Mice

The muskrat is cliaracteristic of the The two species of minids occurring prairie sloughs, marshes, and drainage in the Black Hills are commensals of man, TURNER: MAMMALS OF THE BLACK HILLS 119 and were introduced into North America This rodent was introduced from from the Old World in the mid-eigh- Europe in early times and has since teenth century. spread over most of temperate and tropical North America. I am unfamiliar Rattus norvegicus (Berkenhout) with the history of its dispersal to the Norway Rat Black Hills, but Mus niuscuhis presum- ably was introduced in the plains area Mas norvegicus Berkenhout, 1769, Outlines of throughout the Mississippi and Missouri the natural history of Great Britain and river valleys from river boats to wharves Ireland, 1:5 ( t\'pe locality, England, where forts and it the species was introduced from Asia, prob- of trading posts; thence, ably \'ia Continental Europe, in the early probably spread into the study area in ITOO's). company with early military expeditions Rattus norvegicus—Hollister, 1916, Proc. Biol. and the establishment of settlements. Soc. Washington, 29:126, 6 June. Hayden (1859:710) mentioned that both The Norway rat was introduced to the house mouse and Norway rat were tlie colonies of eastern United States from abundant at all of the fur trading posts the Old World about 1775, escaping from on the Missouri River in the 1850's. ships at various seaports (Silver, 1941: Because two subspecific stocks were I the of its im- 2). do not know history introduced into North America ( Schwarz migration into the Black Hills region, and Schwarz, 1943:65), I choose not to but the species presumably was intro- assign a subspecific identity to speci- duced at rixer towns and forts along the mens from the Black Hills. Missouri in the 1850's and sub- Valley A female taken southeast of Spearfish in a manner similar sequently dispersed in mid-August evidenced six placental to Mus as described in the musculus, scars, whereas another obtained in Wind following account. Cave National Park in late August I know of only two actual specimens showed no signs of reproductive activity. from the study area, but Over and Testicular lengths of two males obtained Churchill (1945:45) indicated that these in June from near Minnekahta were 5. rats were found in the Black occasionally Specimens examined (25).—SOUTH DA- man. Hills, in and around habitations of KOTA: Lawrence County: 1.5 mi S, 0.5 mi E 3800 1; 8 mi SW Lead, 1 (UW). Specimens examined (2).—SOUTH DA- Spearfish, ft, Meade County: Black Hawk, 1 (AMNH). KOTA: Pennington County: Rapid City, 2 4 mi N Rapid Cit>', 3 (SDMT). Pennington County: 12 (10 NRW, 2 Additional /T'corc/.—SOUTH DAKOTA: (AMNH); Rapid City, Glendale, 1 (AMNH). Custer "Black Hills" (Over and Churchill, 1945:45). SDMT); County: Squaw [Grace Coolidge] Creek, 1 (AMNH); 2.5 mi N, 5 mi E Wind Cave, Wind Mus musculus Linnaeus Cave National Park, 3700 ft, 1; Wind Cave Can- Wind House Mouse yon, Wind Cave National Park, 4100 ft, 1; Cave National Park, 1 (USNM). Fall River nat- [Mus] musculus Linnaeus, 1758, Systema County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 1:62 restricted urae, ed. 10, (type locality ft, 2. to Upsala, Sweden, by Thomas, Proc. Zool. Additional record.—SOVTH DAKOTA: Soc. London, p. 147, March 1911). Lawrence County: Spearfish (BB).

The house mouse is a common, often Family ZAPODIDAE—Jumping Mice abundant, rodent that frequently is in- fauna. in the Black jurious to man and to the native Zapodids, represented are salta- It resides in and around human habita- Hills region by one species, in areas of lush tions at lower elevations in the Black torial rodents that abide such as wet meadows or Hills. For example, on 13 June 1967, sev- vegetation is the eral dozen individuals were killed with streamside communities. Aridity factor the distribution .22 dust-shot in grain bins near Minne- major restricting mice on the Northern Great kahta. Fall Ri\er' County. Wild popula- of jumping tions occasionallv are encountered. Plains. 120 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Zapus hudsonius campestris Preble three female jumping mice were trapped west of Hill in a lowland Meadow Jumping Mouse City, marshy bordering Ditch Creek. Small pools of hudsonius N. Zapus campestris Preble, 1899, water were interlaced with hummocks of Amer. Fauna, 15:20, 8 August (type lo- soil that supported of liverworts cality, Bear Lodge Mountains, Crook Co., growths Wyoming). and mosses. Grasses, sedges, and white spruce also were predominant in the Although specimens are absent from immediate area. Additional Meade and Fall River counties, the mea- specimens were obtained under similar conditions dow jumping mouse is common through- near the east end of Deerfield Lake and out the study area. It most frequently along Goldrun Creek, southeast of Deer- occurs in riparian communities along field. Representatives of Z. /k campestris small streams in valley meadows, or in taken near Nemo, Lawrence County, open, moist habitats within coniferous were trapped along the banks of Box- forests that support a low undergrowth elder Creek. In these various habitats, of forbs and grasses. Zapus hudsonius Sorex cinereus, manictilatus, occurs in suitable habitats on the ad- Peromysctis Microtus lon0catidus, M. pennsyJvani- jacent plains, but is most frequent in cus, and Eutamias minimus were trapped the Black Hills up to 6500 feet. This in association with Z. hudsonius. species hibernates in winter, and all Several mice were collected specimens at hand were taken in warmer jumping in 14 months; earliest and latest seasonal rec- non-riparian habitats. A male from mi Hill was at the base ords are 4 June and 27 August, respec- W City trapped of a rock on a wooded tively. large dry slope that was far removed from a source of Jumping mice collected in this study standing water. A male, from the same were trapped along drainage systems, in localitv, was shot in late afternoon as dense grasses and sedges that were in- it ran from beneath the branches of a terspersed with deciduous shrubs and partialh' hollow log that was situated in fallen logs of aspen, birch, willow, white a dry and rocky meadow. Another male spruce, and ponderosa pine. Most of the from near Nemo was trapped on a dry specimens from the Wyoming part of the hillside ponderosa pine trees and Hills, and those individuals from Little among granite boulders. Spearfish, Big Spearfish, and Beaver About 20:30 hrs on 1 Creek canyons were taken under these (MDT) August 1967, I heard in a stand of general conditions. Similarly, five fe- rustling males and three males were trapped in slough grass (Beckmannia syzigachne) the shore of the a large valley of approximately one mile growing along Savoy Lawrence A in length located southwest of Keystone. Reservoir, County. jump- mouse was observed an A stream, with many small tributaries, ing climbing stem of thus rid- flowed through the center of the valley already-inclined grass, and created a semimarsh situation. Lush ing the stem to the ground. Straddling the bent stem and it close to the vegetation was mainly of tall grasses and holding the mouse ro- some very dense patches of clover; de- ground, systematically ciduous shrubs bordered the stream. tated the rachis and removed the seeds. Within about five the seed- Forty-two individuals (22 males and minutes, laden head of the was and 20 females) were taken in the lowland grass bare, the mouse seeds in its habitat of Grizzly and Iron creeks, north- departed carrying which were distended. east of Custer, in mid-June 1967. Ri- cheeks, noticeably parian habitat was composed of grasses, Fourteen of 31 females taken in the spearmint, yellow wood sorrel (Oxalis latter half of June carried 5.8 (4-7) em- stricta), stinging nettle, and ferns; drift- bryos that were 13.1 (3-18) in crown- wood, woodpiles, and shrubs were dis- rump length; four others had 5.5 (4-7) persed throughout. Eleven male and placental scars. Average testicular length TURNER: MAMMALS OF THE BLACK HILLS 121 of 38 males obtained in tliis period was of Krutzsch (1954:441), indicating that 6.7 (4-10). In July, six of 2.3 females Z. h. campestris, originally described showed no signs of reproductive activity, from the Black Hills, contrasts with Z. h. but 11 pregnant females carried 6.1 (5-8) pallidiis to the east in South Dakota in embr\os \\'hich were 12.9 (3-20) in being conspicuously darker and axerag- crown-rump length. The uteri of six ing larger in external and cranial meas- other indixiduals had 3.8 (2-5) placental urements. scars. of testes of 13 Average length Secondary sexual dimorphism is not males obtained in was 6.1 July (4-8). evident in jumping mice from the study Two of six females in Au- captured early area. However, patterns of variation in gust had 5.5 (5-6) placental scars; none size within the Hills are discernible. In- was pregnant. Testicular length of six dividuals from Pennington County are males obtained at this time was 5.7 . (4-7) larger than those from other Black Hills Most females that exhibited placental populations in occipitonasal length, zygo- scars also were lactating. matic length, length of palatine foramen, Three young Zajnis were captured and cranial depth, but are smaller in northeast of Custer in late June. Sub- length of maxillary tooth-row, zygomatic adults, which are brighter in color than breadth, mastoid breadth and width of adults and generally lack a distinctive palatine foramen. These dimensional re- mid-dorsal stripe, are not uncommon lationships are reciprocal when compar- throughout the warm months, until at ing representatives from Lawrence least mid-August. A female li\e-trapped County with those of other areas (Rob- on 25 July 1967 at the last-mentioned ertson, 1971:16-17). Physiographic ir- site gave birth to seven still-born young regularity of the Hills may contribute to during the ensuing night; external meas- this morphological diversity. Coloration urements of these newborn voung are: shows no readily recognizable trends in total length, 38.3 (37-39); tail length, the Black Hills. 10.8 (9-13); and hind foot length, 5.0 Selected external and cranial meas- (5). urements of 20 adult males, followed by The process of annual molt occurs in those of 20 adult females, are (Robert- adults from mid- to late summer. For son, 1971, Table 1): total length, example, five males taken from 27 July 125.3+5.43, 126.2+6.86; hind foot to 2 in Little 29.6 + 29.5+1.64; ear August Spearfish Canyon length, 1.47,+ length, displayed bright-colored tips of new 14.4+1.70, 13.4 1.60; occipitonasal hairs from beneath the old 23.5 + projecting length, 0.54, 23.2+0.46;+ zygomatic dull pelage of the dorsum. Two females length, 9.5+0.25, 9.5 0.28; zygomatic from southwest of Rapid City and two breadth, 11.4+0.35, 11.3+0.39; mastoid males from north of Sundance (all cap- breadth, 10.1+0.38, 10.0+0.45; inter- in were in orbital 4.2 + 0.22, tured mid-August ) completely constriction, 4.3+0.20; new pelage. cranial depth, 9.1+0.16, 9.2+0.27; palate 3.5 + 3.5+0.23; of Robertson (1971) recently analyzed length, 0.24, length 3.6 + 3.5+ variation in populations of Z. hudsonius maxillary tooth-row, 0.17, of foramen, 4.6+ on the Great Plains. Because his studies 0.15; length+ palatine were concurrent with my own, and be- 0.29, 4.4 0.22; width of palatine fora- cause we had a mutual exchange of in- men, 2.2+0.14, 2.1ih0.14. I his results herein. formation, rely upon Specimens examined (207).—SOUTH DA- Jumping mice exhibit clinal variation on KOTA: Lawrence CounUj: Big Spearfish Can- 9 mi 3 mi 5000 ft, the plains northwestward from eastern yon, S, W Spearfish, 9; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, Kansas, becoming progressively darker 5800 ft, 14; 2 mi S Tinton, 6100 ft, 7; 3 mi in coloration and tending toward longer, W Nemo, 4800 ft, 1; 2 mi W Nemo, 4700 ft, skulls. Robertson's narrower, deeper 9; Nemo, 4700 ft, 1. Pcunin^fon Conniij: 1 Beaver findings (1971:20-21) agree with those mi S, 8.5 mi W Rapid City, 2; Creek, 122 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

4 mi N, 10.5 mi W Deerfield, 6400 ft, 10; lacking from Fall River and Weston Beaver Creek, 3 mi N, 9 mi W Deerfield, 6400 counties. Goldrun Creek, 1 mi S, 2 mi E Deer- ft, 1; Erethizon dorsatum hruneri differs field, 6400 ft, 3; Rapid Creek, 2 mi W Pactola, T. from E. d. which occurs to 4800 ft, 3 (UMMZ); Castle Rock, R. 2 E, epixanthum, the in in 1 N, 6500 ft, 3 (UMMZ); Ditch Creek, 14 west Wyoming, ha\'ing smaller Ditch 13 mi W Hill City, 6400 ft, 10; Creek, external and cranial dimensions, arched mi W Hill City, 6400 ft, 4; Nelson's Place, frontals, and auditory bullae that are less 3 mi SE Hill Cit\% 6 ( UMMZ ) ; Palmer Gulch, inflated 4 mi SE Hill City, 6 (UMMZ); mispecified (Long, 1965:669). locality, 1 (UMMZ). Citster County: 3.5 mi Porcupines usually were encountered Palmer S, 0.5 mi W Kevstone, 5000 ft, 8; along the shoulders of highways in eve- Gulch, 8 mi SE Hill City, 9 (FMNH); 5.75 ning or early morning, and many speci- mi N, 5.75 mi E Custer, 5220 ft, 42; Custer, 5 mens in collections were taken as road- (2 AMNH, 3 USNM); Wind Cave National kills. Due to inflicted Park, 1 (WCNP); Game Ranch, Cold Spring heavy damage by Creek, Wind Cave National Park, 2 (UMMZ); this species upon potentially harvestable Bea\'er Creek Wind Cave National Canvon, timber, particularly young pine trees, the Park, 4200 ft, 9 (I'UMMZ). U. S. Forest Service maintains a porcu- WYOMING: Crook County: Bear Lodge pine control program in the Hills. Trees Mountains, 6 ( USNM ) ; Bear Lodge Mountains, 6.5 mi SSE Alva, 1 (UMMZ); Bear Lodge with tops and branches barked indicate Mountains, Warren Peak Lookout, 6000 ft, 2 the abundance and widespread distribu- Dexils flood Belle (USNM); Tower, plain tion of Erethizon in the study area. Fourche River, 3350 ft, 1 (USNM); 15 mi N Sixteen indi^'iduals were collected Sundance, 2; 3 mi NW Sundance, 5900 ft, 19; Sundance, 3 (USNM). Weston County: 1.5 along Little Spearfish Canyon in late mi E 6150 7. Buckhorn, ft, July and early August of 1967, generally Additional recorJ.s.—SOUTH DAKOTA: along roadways at dusk. On t\\'o occa- Latcience County: Spearfish (BB). Penning- sions, juvenile males were taken in com- ton County: Rochford (BB). Custer County: with adult males. Bull Springs (Bole and Moulthrop, 1942:167). pany large One procu- pine was shot as it persistently chewed Family ERETHIZONIDAE— a hole in a camper's tent; bacon grease Porcupines had been spilled on the tent and even repeated attacks on the porcupine with one with one of Only genus, species, a club could not drive the rodent awa\'. this rodent occurs in the large, quilled Another indi\idual was shot at mid-day Black Hills region. as it roosted in the top of a pine at the Erethizon dorsatum bruneri Swenk bottom of Big Spearfish Canyon. bluffs are used as den sites Porcupine Rocky by Erethizon; dens are evident from ac- Erethizon epixanthum hruneri Swenk, 1916, cumulations of feces and discarded quills. Univ. Nebraska Studies, 16:117, 21 Novem- Porcupine scats are in the form of pellets, ber (type locality, 3 mi E Mitchell, Scott's which are often interconnected Bluff Co., Nebraska). by strands of Fre- Erethizon dorsatum hrw^eri—Anderson, 1947, undigested vegetation. Bull. Nat. Mus. Canada, 102:173, 24 Janu- quently dens of Erethizon are associated ary. with those of Marmota faviventris. Erethizon dorsatum occurs on the A female taken near Rockerville on Great Plains and adjacent montane areas 29 March 1946 contained one embryo. at sites where trees are available locally. Females obtained in July and August In the Black Hills, porcupines have a each showed a single placental scar in the wide altitudinal distribution, occurring uterine horns. A lactating female was from the boreal cap, down through the collected northeast of Custer on 17 June transition zone, and out onto the sur- 1967, and juveniles were common from rounding prairie. This rodent is active July through September. The average the year around. Although common in length of testes of four males taken in the Hills, specimens of Erethizon are August was 44.2 (40-57). Ju\enile pelage TURNER: MAMMALS OF THE BLACK HILLS 123 consists of fine soft blackish underfur sult, the gray wolf, black bear, grizzly and long yellowish guard hairs, as com- bear, mountain lion and lynx either no pared to the rather coarse underfur and longer occur in the Hills, or are rare. whitish guard hairs of adults. The quills Coyotes, badgers, and bobcats also must are less resilient than tliose of adults, endure the attempts of humans to deci- and also possess a yellowish tinge. mate their populations. Specimens examined (52).—SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- Family CANIDAE—Coyotes, mi 4200 5900 yon, 3 W Spearfish, ft, 1; Tinton, Wolves, and Foxes ft, 1; 2 mi S Tinton, 6100 ft, 1; Little Spear- fish Canyon, Roughlock Falls, 9 mi W Lead, This family of doglike mammals is 1 mi 5400 ft, 1; Little Spearfish Canyon, S, represented in the area under study by 9 mi W Lead, 5400 ft, 5; Little Spearfish Can- two genera and three species. Two addi- yon, 2 mi S, 10 mi W Lead, 5800 ft, 7; Little tional canids, the swift fox and gray fox, Spearfish Canyon, 6.2 mi SW Lead, 6000 ft, occur in the and oc- 1; Little Spearfish Canyon, 12 mi S, 14 mi W general vicinity may Lead, 6200 ft, 2; Bogus Jim Creek, 1 (SDMT). casionally intrude into the Black Hills. Meade County: Vanocker Canyon, 3 mi S, 0.5 mi W Sturgis, 4400 ft, 1. Pennington Canis latrans latrans Say County: 1.5 mi N Silver City, 1 (SDMT); 0.4 mi SW Silver City, 1 (SDMT); Rapid City, Coyote 1 (NRW); Bea\'er Creek, 3 mi N, 9 mi W Cams latrans Say, 1823, in Long, Accoimt of an Deerfield, 6500 ft, 1; 1 mi NW Deerfield, 1; expedition from Pittsburgh to the Rocky Deerfield, 1 (SDMT); Spring Creek Canyon, Mountains . . ., 1:168 (footnote), (type 4 mi ENE Rockerville, 3600 ft, 2 (UMMZ); locality. Engineer Cantonment [approxi- Ditch Creek, 14 mi W Hill City, 6400 ft, 1; mately 2 mi E Ft. Calhoun], Washington 19 mi W, 3 mi S Hill City, 1; Palmer Gulch, Co., Nebraska). 4 mi SE Hill City, 5300 ft, 1 (UMMZ); un- specified locality, 2 (SDSU). Custer County: The coyote is known from much of 2 mi N, 22 mi W Custer, 5200 RoJjy Canyon, North America, and is well distributed ft, 1; 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; the Black Hills. At least 200 Shirttail Canyon, 2 mi S Wind Cave, Wind throughout the Sheid- Cave National Park, 1 (WCNP); Dewey, 11 individuals were poisoned on (USNM). lev Cattle Companv range near Rapid WYOMING: Crook 15 mi N Sun- County: city in 1894 (Bailey, 1907:11), 1165 dance, 1; 15 mi ENE Sundance, 1; Sundance, coyotes were killed in the Bear Lodge Bear Lodge Mountains, 6000 ft, 1 (USNM). Mountains in 1907 1908:6), and Weston County: 1.5 mi E Buckhorn, 6150 ft, 1. (Bailey, Additional records.—SOUTH DAKOTA: reports of District Rangers, Black Hills Custer County: "vicinity of Otis" (Stebler, National Forest (U. S. Biological Survey 19.39:389). WYOMING: Weston County: files), indicate that approximately 2680 "east of Newcastle" (Stebler, 1939:389). coyotes were killed in that area in the period from 1926 to 1935. From 1962 ORDER CARiNIVORA—Carnivores to 1968, bounties on 19.30 coyotes were A rather diverse group of mammals paid by the counties that comprise the to the robust Hills rec- from the diminutive ermine South Dakotan portion of the ( grizzly bear, comprise the carnivorous ords from South Dakota Dept. Game, fauna (nine genera and 15 species) in Fish and Parks), and extensive use of the Black Hills within historic time. The 1080 poison (sodium monoHuoracetate) statuses of an additional seven species has further reduced the population in (si.x genera) in the Hills region remain the eastern and southern sections of the uncertain. Black Hills (A. H. Richardson, pers. Representatives of all five families of com.). In spite of these attempted con- Carnivora that occur in North America trol measures, Canis latrans seems to be either are indigenous to the Black Hills increasing in abundance in the northern or recently have been extirpated by man. Hills, especially along the South Dakota- Large carni\'ores tend to interfere with W\'oming border (A. W. Jones, pers.

as a re- com. . man's pastoral enterprises, and ) 124 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Compared to Canis latrans Jestes, be found in the subsequent inspection which occurs to the west in Wyoming, along the route of the chase. C. /. latrans is slightly smaller externally Reproductive data applicable to the and cranially, and the upper parts are Black Hills coyote population were ob- much paler in color (Long, 1965:674). tained from the U. S. Biological Survey files. females taken near Custer Evidently, coyotes increasingly in- Two vaded the Black Hills as gray wolves in March 1944 carried four and sexen nine females ob- were extirpated by man, and became embryos, respectively; much more abundant as the latter species tained in the Hills in late April and had an of 6.4 was reduced in numbers. Grinnell ( 1875: early May average (4-7) each. females 79-80) stated: "The coyote was found embryos Two adult (one with se\'en and the other in considerable numbers on the plains, suckling pups with four were in and was especially abundant among the embryos) captured elevated table-lands that were crossed Wind Cave National Park late in April 1918 C. Beach. just before reaching the Black Hills. by Troy Thus, coyotes the After penetrating into the hills proper, in the Black Hills, as elsewhere on northern breed in Feb- however, I did not see a single specimen plains, apparently until I left them for the Big Cheyenne ruary and March, bearing litters in April [Cheyenne River], when I again noticed and May (Mengel, 1971:325). coyotes in numbers. In the Black Hills, Stomachs of five coyotes, trapped in this species would seem to be replaced Wind Cave National Park in early June by the preceding [gray wolf]." In 1894, 1924, contained the remains of domestic W. W. Granger indirectlv referred to a chicken, domestic lamb, and cottontail change in relative abundance of the rabbit (Sylvilagus sp.). two when he noted that species gray Specimens examined (19).—SOUTH DA- wolves were "not uncommon," and coy- KOTA: Pennington County: 4 mi S, 14 mi W otes were "common" in the Black Hills Hill City, 2. Custer Countij: 18 mi NE Dewev, 4 (USNM); 12 mi NE Dewev, 1 (USNM); (J. A. Allen, 1895a: 274). Coyotes con- 4 mi N 1 5 mi NW Wind tinued to increase in numbers in the Dewey, (USNM); Cave, 1 (WCNP); Wind Cave Canyon, Wind Hills after the turn of the century, Cave National Park, 1 (WCNP); Shirttail Can- whereas the gray wolf was eliminated yon, Wind Cave National Park, 2 (WCNP); 3 mi SE 1 Fall River shortly thereafter. Conversely, the de- Wind Cave, (WCNP). Count!/: 5 mi W Minnekahta, 1. cline in ungulate populations, important WYOMING: Crook County: Bear Lodge as a source of food for and in- coyotes, Mountains, 2 (USNM); Grand Canyon, Sun- creased conflict with the agricultural pur- dance, 1 (USNM); Rattlesnake Canyon, Sun- suits of man, caused a reduction in the dance, 1 (USNM). number of C. latrans on the plains. UNSPECIFIED LOCALITY: Black Hills, 1 (SDSU). I often have observed C latram near Additional records card files unless dusk and after dawn in the (USES shortly otherwise noted).—SOUTH DAKOTA: Law- various towns in Wind Cave prairie dog rence County: Spearfish; Lead. Meade County: National Park. On 25 July 1968, I Piedmont. Pennington County: Rochford; watched a large adult "mousing" along Deerfield; 17 mi NW Custer. Custer County: Bull 14 mi Custer 1943: Highland Creek for several hours in mid- Springs, NW (Lamster, 1); Custer; 9 mi W Pringle; Pringle; Dewey; A litter was raised in Beaver morning. 18 1 mi W Custer; unspecified locality, (Jack-' Creek that summer and the Canyon pups son, 1951:262). Fall River County: Hot were vocally active each evening. On 27 Springs; Minnekahta. August 1968, an adult coyote and three young were observed pursuing a doe Canis lupus irremotus Goldman pronghorn in Shirttail Canyon. The fate Gray Wolf of the pronghorn remains unknown as Canis lupus irremotus Goldman, 1937, Jour. all animals from (quickly passed view; Nhmun., 18:41, 14 February (type locality, however, no evidence of its capture could Red Lodge, Carbon Co., Montana). TURNER: MAMMALS OF THE BLACK HILLS 125

The gray \vo\i formerly occurred on ihc range of this company in 1896 throughout much of the United States. (Bailey, 1907:11). Near Newcastle, Historical reports indicate that Conis wolves were still numerous enough to be lupus pre\iously was abundant and destructive to li\estock in 1905 and 1906 widely distributed on the Great Plains (Bailey, 1907:10). Evidently, the gray and in the Black Hills. Extirpation of the wolf was already becoming more scarce vast herds of bison destroyed the main at this time, whereas the coyote was food source of the \\'olf and this, along becoming more abundant in the Black with trapping and poisoning of wolves Hills (see Allen, 1895a: 274). for of for and protection livestock, fur, Decimation of C. lupus in the Hills for led to the elimina- bounties, finally continued during the early part of the tion of this from areas of species major twentieth century (U. S. Biological Sur- its former range. . vey files, unless otherwise noted ) About Conis lupus irremottis differs from 925 gray wolves were obtained by the C /. nuhilus to the south and east in U. S. Forest Service in the Bear Lodge being larger, paler, and narrower in the Mountains and surrounding portions of frontal region, especially relative to the Crook County, Wyoming, in 1907 width of the rostrum (Long, 1965:677). (Bailey, 1908:6). Bounties were paid in Goldman (1944:448-449) noted that 1911 on 55 wolves by counties that com- specimens from the Black Hills are vari- prise the South Dakotan section of the able and grade toward nubilus. Hills (records from South Dakota Dept. and wolves Grinnell (1875:79) wrote of this spe- Game, Fish, Parks). Eight were in and five cies as follows: "I found the gray wolf destroyed 1915, more one of the most common animals in the were killed in 1916. A pregnant female that carried four was taken 18 Black Hills, and hardly a day passed embryos mi NE on 25 March 1917. A without my seeing several individuals of Dewey male was killed 15 mi SE on 10 this species. They were generally ob- Dewey November 1920. Three indi\'iduals were served singly or by twos and threes, shot on 23 December and two were sneaking along the mountain sides or 1926, taken on 11 1928. crossing the narrow valleys." He indi- January cated that in the Black Hills, C. latrans One of the famous American rene- (see account of this species) was re- gades was the "Custer Wolf" which placed by C. lupns. Dodge (1876:133) ranged in the vicinity of Custer for a

noted : "The wolves are of enormous size, period of seven years (Young, 1946a: 68, a very dark gray in color, and in consid- photo opposite). This large male had a erable numbers." From the spring of known range of 40 by 60 miles (Seton, 1895 to the autumn of 1897, about 500 1929a: 257) and was credited with killing gray wolves were killed on the range of $25,000 worth of livestock during this the Ames Cattle Company (54.35 square period. It avoided capture despite a miles) in northeastern Crook Count}', bounty of $500 off^ered for its scalp. A T. Federal H. P. concen- Wyoming (Long, 1965:676). J. Craig, Hunter, Williams, manager of Western Ranches Limited, trated entirely upon the "Custer Wolf" Belle Foiu"che, South Dakota, filed a re- from March to October 1920 before fi- port with the U. S. Biological Survey in nally succeeding in killing it (Young, 1896, indicating that C. lupus was re- 1944:277). The last specimen from the sponsible for a three percent loss of Black Hills known to me is one reported calves, one percent loss of adult cattle, by the Distiict Forester in Harney Na- and five percent loss of colts annually tional Forest, 30 January 1934 (U. S.

files . 4 (Young, 1946a: 122). G. E. Lemmon, Biological Survey ) On July 1967, manager of the Sheidley Cattle Com- I found a mandible of C. lupus in a pany, Rapid City, South Dakota noted woodrat nest at the rear of Davenport that 80 to 100 gray wolves were poisoned Cave, southwest of Sturgis. Currently, 126 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY the former range of C. lupus in the Black eastern sections (see additional records Hills has been pre-empted by Canis for specimens that recently have come latrans. to the attention of these men). A red fox was taken Specimens examined (3).—SOUTH DA- recently along KOTA: Meade County: Da\'enport Cave, 3 mi the southern periphery of the Black Hills S, 1.5 mi W Sturgis, 4400 ft, 1. Custer County: at a locality 10 mi N Rumford, Fall Ri\'er 12 mi NE 1 Dewey, (USNM). Gounty (Jones and Henderson, 1963: WYOMING: Crook County: Sand Creek 283). 'From 1962 to 1968, bounties on Canyon, 1 (USNM). Additional records.—SOUTH DAKOTA: 1079 V. vulpes have been paid by those Custer County: 18 mi NE Dewey (USES counties that comprise the South Da- files); Dewey (Goldman, 1944:445). Fall kotan of the Hills portion Black ( records River County: 15 mi SE Dewey (USES card from South Dakota Dept. Game, Fish, files). WYOMING: Crook County: northeast- and Parks . ern portion of county (Long, 1965:676). ) Vulpes vulpes regalis diflFers from V. V. macroura of western in Vulpes vulpes regalis Merriam Wyoming and more Red Fox being larger golden-reddish instead of yellowish in color (Long, Proc. Vulpes regalis Merriam, 1900, Washing- 1965:679). As other recent authors, I ton Acad. 28 December Sci., 2:672, (type follow Ghurcher 1959 in the ( ) using spe- locality. Elk River, Sherburne Co., Minne- cific name vulpes sota ) . Vulpes (Linnaeus) Vulpes fulva regalis—E. Bailey, 1929, Jour. for American red foxes. Mamm., 10:157, 9 May. Specimens examined (2).—SOUTH DA- The red fox is widely distributed and KOTA: Custer County: Custer, 1 (USNM); Elk Mountain, 1 (USNM). relatively common in the eastern Great Additional records (A. H. Richardson, L. Plains, and occurs sparingly in suitable Petersen, and E. L. Woods, pers. com.).— habitat in the eastern and southern por- SOUTH DAKOTA: Lawrence County: near tions of the Black Hills. Grinnell (1875: Nemo, 1. Pennington County: 7 mi NW Rapid 1. Custer 2 mi S 2 96) observed three red foxes, an old City, County: Custer, 1; mi W Hermosa, 1. Fall River 2 mi female and two young that inhabited County: N Hot Springs, 1; 2 mi S Hot Springs, 1. a burrow on the edge of a prairie dog town the northern of the along fringe Family URSIDAE—Bears Black Hills in 1874. Bailey (1888:432) The omnivorous ursids are witnessed "a few tracks" of this species large rep- resented in the Hills in the snow in October 1887, and on 20 Black by one genus and two July of the following year obtained a species. female near Custer, noting a field report Ursus americanus americanus Pallas to the U. S. Biological Survey that this species was "common" in that vicinity. Black Bear A male was taken Merritt near by Gary Ursus americanus Pallas, 1780, . . . Spicilegia

Elk Mountain on 16 October 1903. zoologica . . ., fasc. 14.5 (type locality, eastern North x\merica). Game Biologists A. H. Richardson and L. Petersen, and Game Warden E. L. Although I have examined but a Woods, indicated to me that Vulpes single specimen of this species, historical vulpes was relatively unknown in the accounts as well as recent newspaper Black Hills five to 10 years ago, but that accounts and reports by National Forest since 1965 it has increased noticeably in Service personnel, indicate the presence the region (pers. com.). These natural- of this bear in the Black Hills. ists felt that the coyote population has Dodge (1876:132) wrote of bears in decreased considerably in some areas of the Black Hills as follows: "The country the Black Hills because of extensive use in this part of the Hills [along Rapid and of 1080 poison, and that the red fox is Boxelder Greeks in Pennington Gounty] the replacing coyote in the southern and is full of bear sign. In some places almost TURNER: MAMMALS OF THE BLACK HILLS 127

a quarter of an acre will be rooted up as Ursus aretos horribilis Ord if small thickets of by hogs; berry-bear- Grizzly Bear ing bushes are torn and broken; ant-hills are dug into and huge logs turned over C7r,v(/.v hornhilis Ord, 1815, in Guthrie, a new by this omnivorous monster in search of geograpliical, historical, and commercial grammar . . ., ed. 2, 2:291, 299 (type lo- his food." He (1876:133) continued, cality, north side of Missouri River, near . . seven or bears were killed eight by moutli of Wolf Creek, Roosevelt Co., Mon-

our . . . . whole party from the little tana ) black to the mammoth grizzly." Indians Ursus aretos horribilis—Erdbrink, 1953, A re- vie\\' of fossil and recent of told Hoffman (1877:97) that the only bears the Old World. . . . de 30 March. place the black bear was encountered Jan Lange, p. 339, Ursus rogersi bisonophagits Merriam, 1918, N. was in the Black Hills and in the Big- Amer. Fauna, 41:66, 9 February (type lo- horn country. men- Bailey (1888:432) cality. Bear Lodge Mountains, Sundance tioned that three kinds of bears (black, National Forest, Black Hills, Crook Co., brown, and silvertip) "were said to oc- Wyoming ) . cur" in the Hills. In discussing the Black Hills in 1851, On 2 a 160 September 1968, pound a trapper indicated that "Grizzly bears ! domestic ewe, with its throat slashed and are found there sometimes in bands like Hesh eaten from its and back, shoulders, buffalo. . . ." (Culbertson, 1952:90). was found in Swede hindquarters, Gulch, Although not quite as enthusiastic, Grin- on the Wolfe 2 mi N and Philip ranch, nell (1875:81), Dodge (1876:132-1.33), 5 mi W Rochford, Pennington County and Bailey (1888:432) each alluded to 3 (Rapid City Journal, September 1968, the presence of large numbers of grizzly p. 2). On the stream bank beside the bears in the Black Hills in the late 1800's. ewe was the distinctive of a bear. print The four specimens examined all were In November of the same a 300- year taken in the study area between 1855 adult male black bear was shot pound and 1887. On 7 August 1874, General west of Rochford, between Black Fox Custer and Colonel Ludlow shot a well- and Crook's Tower. Campground scarred old male near Nahant (about Mr. Raymond Nilles, Forest Ranger three miles south of the present town of of the Sundance District, told me of find- Dumont, Lawrence County, South Da- ing huge, deep claw marks on a tree kota); soon after, other members of the along Hershey Creek in the northern expedition killed a cinnamon-colored Hills in 1966. in the area 1875: The claw marks began grizzly same ( Winchell, about six feet from the base of the tree 50; O'Hara, 1929:251-252). The male and extended to the ground; Mr. Nilles was a deep, glossy black, except for a and others credit these markings to the sprinkling of dark gray hairs on the activity of a bear. Other recent reliable head, lower parts of the shoulders, and sightings of black bear in the Black thighs (Grinnell, loc. cit.). Two Indian Hills are as follow: Big Spearfish Can- scouts killed an old female and two cubs yon; along Cold Creek, in Lawrence a few days later much farther eastward County; near Redfern; in Beaver Creek (along Boxelder Creek, Lawrence Co., Valley, in Pennington County; and at South Dakota). "The cubs were about Camp Mallo in Weston County. It half grown, and with the mother, were should be noted that captive black bears of a yellowish clay color. The inner half are tourist attractions in the Black Hills of each hair was deep black, but the a reddish- region, and some of these may occa- outer extremity was bright This them a curious mottled sionally escape. yellow. gave appearance and induced many of those examined 1 . —SOUTH DA- Specimen ( ) who saw them to consider them a differ- KOTA: Pennington Cotintij: west of Rochford, ent from the one killed Gen- 1 (mounted and on display at Wall Drug species by Store, Wall, South Dakota). eral Custer" (Grinnell, loc. cit.). 128 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

The foregoing account points out the its flesh eaten by humans; it provides extreme amount of individual variation much recreation as a hunted fur-bearer, found among this species, and the taxo- and accounts for substantial loss of cer- nomic dilemma thus implied. At the tain crops to farmers annually. The spe- moment, there are about 96 named kinds cies probably is commoner in the region of grizzly bears, most of which can be now than in the recent past. attributed to the work of C. Hart Mer- lotor hirtus riam. iMerriam (1918:66) described Ur- Procyon Nelson and Goldman siis rogersi hisonophagus from the Bear Lodge Mountains, and at the same time Raccoon included the Black Hills (1918:93) Procyon lotor hirtus Nelson and Goldsmith, within the range of Ursus absarokos. I 1930, Jour. Mamm., 11:455, 11 November follow Erdbrink (1953:339), who re- (type locality. Elk River, Sherburne Co.,

Minnesota . garded the brown bear of the Palearctic ) and Nearctic regions as being conspe- Procyon lotor is common and wide- cific and therefore employed the name spread in suitable habitat throughout Ursus arctos horrihiJis. Professor E. Ray- North America. Although few actual mond Hall, of The Universitv of Kansas, specimens have been taken, the species currently is reviewing the taxonomic is common in the Black Hills. Usually status of the grizzly bear. The following the raccoon is found along streams, selected cranial measurements of grizzlys around lakes and marshes, and is a fre- from the Hills were taken by Professor quent nocturnal visitor to area camp- Hall, and are the average (extremes) of grounds. three females and a male of U. r. (type Procyon lotor evidently is a recent all animals: hisonophagus) , fairly young addition to the Black Hills mammalian 281.3 occipitonasal length, (255.0-322.0); fauna. Hayden (1859:706, 1862:143) 166.5 inter- palatal length, (157.0-180.0); noted that this species was seldom seen orbital 70.5 breadth, (63.2-79.4); length beyond the frontier in the mid-nineteenth of 72.2 of P4-M2, (68.0-76.2); length century, and indicated that, although a M2, 36.5 (33.7-38.4); breadth of M2, 18.1 few had been obser\ed in the White (17.2-19.7). River Valley, the raccoon seldom passed Green (1961:147, 1962:3) recently lo- up the Missouri River above latitude 42°. cated a partially fossilized arthritic skele- Grinnell (1875) and Dodge (1876) did ton of a grizzly in a cave near Hot not include the raccoon in their respec- Springs, Fall River Co., South Dakota. tive lists of mammals encountered by It should be noted that all reports of early expeditions to the Black Hills re- in Hills bears the Black region since the gion. turn of the century have been credited A careful search of the mud banks of to U. americanus rather than U. arctos; almost any riparian community in the even rare in evidently, though occurence, Hills will reveal the presence of tracks the former has the latter in replaced of P. lotor. While camping in Little recent historic time. Spearfish Ganyon in early August 1967,

examined 4 . — : Specimens ( ) WYOMING my cooking supplies were raided each Crook Bear 1 County: Lodge Mountains, night by a large male of this species. A (USNM); Sundance National 3 Forest, steel anchored a (USNM). huge trap, by heavy chain and baited with carcass of a chip- munk, finally captured the raccoon, but Family PROCYONIDAE—Raccoons it escaped, dragging the trap behind. Only one procyonid species, the rac- One week later a local fisherman found coon, is distributed in the Black Hills. the animal, with the trap and chain still The raccoon is an economically impor- attached, drowned in Little Spearfish tant mammal. Its pelt is marketable and Greek. An adult Procyon and four ju- TURNER: MAMMALS OF THE BLACK HILLS 129

\xMiilcs were observed in the area of feet. This small weasel is rather un- Uoughlock Falls on 7 August of the same common in collections due to its seclusive \ear. Another indi\idual was a nightly habits and is known only from Penning- visitor at the Boxelder Creek Camp- ton and Custer counties in the South of the but from ground, just \\ est of Nemo, in late June Dakotan portion Hills, 1967. Specimens of this species obtained both counties of the Wyoming section. in the vicinity of Hill City were taken Hall (1951:164-165) first reported this along a small tributary in a spruce can- species from the study area. data the er- yon. Ecological concerning mine in the Black Hills are rather Specimens examined (4).—SOUTH DA- the winter months of KOTA: Lawrence County: 2 mi S, 10 nii W meager. During 1. 4 mi SE indi- Lead, 5800 ft, Pennington County: 1945-1946, J. A. King captured five S 1 Hill City, 5300 ft, 1 (UMMZ); Hill City, viduals (two males and tliree females) (U\LMZ). Custer County: Wind Cave Na- in the central Hills in box traps. The tional Park 1 (WCNP). in and Additional records.—SOUTH DAKOTA: traps were placed aspen groves Custer County: Custer State Park (USES card along marshes bordering small tribu- files). WYONHNG: Crook County: New taries that flowed through canyons domi- Haven 1950:38). (Goldman, nated by spruce. One female that was kept as a pet by King, carried four em- Family MUSTELIDAE—Weasels, bryos when she died on 30 March 1946. Skunks, and Allies Two females were obtained in mid-July Three genera and six species of of 1965 on a huge dirt slide, overgrown weasel-like mammals inhabit the Black with grass and small aspen, situated be- Hills; the occurrence of four additional tween two high, rocky outcrops east of five in the area One had raised genera ( species ) general Four Corners, Wyoming. remains uncertain. Members of this a fitter shortly before capture; three old family mainly are terrestrial, but two placental scars were present and mam- (the badger and black-footed ferret) mary tissue was receding. The other was are semifossorial in habit, and one (the a subadult, possibly a young of the first mink) is equally at home on land or in female. Coats (1945:10) captured 16 water. Breeding female mustelids gener- ermine one winter in the vicinity of Hill ally exhibit induced ovulation, delayed City. implantation, and are monestrous. Both Individuals taken in the warmer sexes possess anal scent glands, of which months (June and July) were in brown those of weasels, mink and skunks are summer pelage and those obtained in the most noisome to humans. colder months (November to January) were in white winter pelage. Three er- Mustela erminea muricus (Bangs) mines that died in March were molting Ermine from winter to summer pelage; two of as Putorius (Arctogale) muricus Bangs, 1899, these had been kept pets by King Proc. New England Zool. Club, 1:71, 31 since mid-January, and were in the 21st 7500 El Do- July (type locality. Echo, ft, day (UMMZ 90189) and 37th day rado Co., California). (UMMZ 90190) of molt, respectively, at Mustela erminea murica— Univ. of Hall, 1945, time of death. Kansas Publ, Mus. Nat. Hist., 25:84, 27 examined —SOUTH DA- February. Specimens (12). KOTA: Pennington County: Palmer Gulch, Mustela erminea muricus is distrib- 3 mi SE Hill City, 5300 ft, 2 (UMMZ); Pflan- uted throughout the northwestern quar- der's Ranch, 3 mi SSE Hill City, 5300 ft, 1 4 mi SE Hill ter of the United States; in the Black (UMMZ); Nelson's Place, City, of 5300 ft, 1 (UMMZ); Spring Creek, south Hills it is most common in lower spruce- Rapid Citv, 2 (MHM); Spring Creek, 2 mi W streamside coniferous aspen parklands, Ore%ille, 5500 ft, 1 (UMMZ). Custer County: wood- forest, and grassy to semi-swampy 0.5 mi E Sylvan Lake, 6250 ft, 1 (UMMZ). 3 mi lands, usually at altitudes above 5000 WYOMING: Crook County: NW 130 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

4 mi in Sundance, 5900 ft, 1. Weston County: E around headquarters buildings Wind Four Corners, 6000 ft, 2. Cave National Park, but no specimens UNSPECIFIED LOCALITY: Black Hills, were obtained. J. A. King captured a 1 (SDSU). male in an aspen thicket southeast of Hill in of this indi- Mustek frenata alleni (Merriam) City January 1946; vidual was retained in until 9 Long-tailed Weasel capti\ity April. King noted that molt from white Ptitorius alleni N. Amer. Fauna, Merriam, 1896, winter pelage to brown summer pelage 11:24, 30 June (type locality, Black Hills, commenced about 4 February; thus, Custer, Custer Co., South Dakota). when the male (UMMZ 90209) died, it Mustela frenata alleni—Hall, 1936, Carnegie was in the 64th of Inst. Washington Publ., 473:106, 20 Novem- day pelage replace- ber. ment and long white guard hairs were still present over shoulders and neck. The long-tailed weasel occurs loc. cit. noted that the Novem- throughout the United States. In the Bailey ( ) ber-taken individual ". . . is Black Hills it inhabits brushy areas, just chang- from the brown coat to the white, woodlands, and rock piles, usually above ing and is a little more than half white. The 5000 feet and near a source of permanent brown hairs come out very easily; the water. The species e\'idently is rather white hairs are firm." He also witnessed uncommon in the Hills, being presently a few tracks creeks and ventured kno\\aT[ only from Pennington, Custer, along weasels were com- and Crook counties. This mustelid was that long-tailed quite mon in the area. first recorded from the study area by general examined (13).—SOUTH DA- Vernon Bailey (1888:433). Specimens KOTA: Pennington County: Hill City, 1 As presently understood, Mustela (AMNH); Palmer Gulch, 3 mi SE Hill City, alleni is confined to the Black frenata 1 (UMMZ); 20 mi N Elk Mountain, 6000 ft, Hills of South Dakota and Wyoming. 1 (USNM). Custer County: Palmer Gulch, 8 Hill 5300 4 Bull Hall (1951:276) recorded alleni from mi SE City, ft, (FMNH); 2 mi N, 9 mi W Custer, 1 (UMMZ); Mitchell, Scott's Bluff Co., Nebraska, but Springs, Custer, 3 (2 AMNH, 1 USNM). Jones (1964:270) subsequently assigned WYOMING: Crook County: Bear Lodge these to the specimens subspecies longi- Mountains, 3 mi N Sundance, 5500 ft, 1 caucla. Specimens from northwestern (USNM); Sundance, 1 (USNM). Nebraska, and one from Rapid City, Mustela nigripes (Audubon and Pennington Co., South Dakota, undoubt- Bachman ) edly represent intergrades between alleni Black-footed Ferret and longicauda. Putorius Audubon and Bachman, 1851, Mustela frenata alleni difters from nigripes The of North Amer- to the viviparous quadrupeds M. f. longicauda, which occurs ica, 2:297 (type locality. Ft. Laramie, east and south of the Black Hills, north, Goshen Co., Wyoming, according to Hay- in possessing smaller cranial and external den, Trans. Amer. Philos. Soc, 12 (n. s. ): dimensions. Mustela frenata nevadensis, 138, 1863). known from the west in Wyoming, aver- Mustela nigripes—MiWer, 1912, Bull. U. S. Nat. Mus., 79:102, 31 December. ages slightly larger in size and is a darker list brown in summer pelage than is M. f. Presently on the of endangered alleni (Long, 1965:693). species, the black-footed ferret is among carnivores. Ecological data are essentially lack- the rarest of North American short ing for this species in the Black Hills. In This mustelid is characteristic of November 1887, Vernon Bailey caught a and mid-grass prairies, and its geographic with that long-tailed weasel in a trap baited with range is generally coincident a prairie dog and set on a creek bank of the sciurid genus Cijnomys. Although of food near Rapid City. During the summer ferrets utilize alternate sources of 1968, individuals were observed along (Cahalane, 1954:423; Hillman, 1968:6; stream drainages, brushy ravines, and Henderson et al, 1969:22-24), prairie TURNER: MAMMALS OF THE BLACK HILLS 131 dogs are its primary prc\^ and burrow dog town 2 mi W Midland, Haakon Co., systems of the latter species are inhabited South Dakota, by Mr. George Barnes by ferrets. early in August 1953 (Garst, loc. cit.). Mustela niiiripes currently seems to Subsequently, a Park Naturalist noted a be holding its own in western South ferret in the vicinity of Norbeck Dam Dakota, but changing land use and man's on 27 August 1956. The last actual sight- effort to control prairie dog populations ings of this species in the Park were in through destruction of thousands of dog 1957 (Wind Cave National Park files). towns have adversely affected the distri- When excavating prairie dog burrows, bution and abundance of black-footed ferrets form a characteristic trench-like ferrets throughout its range. For ex- structure from three to five inches in ample, widespread use of compound 1080 width and from one to nine feet in length to eradicate Cijnomys is a potential (Henderson et al, 1969:16). In 1969, threat to the survival of ferrets. Hillman the diagnostic trench of a black-footed (1968:11-12) has shown that consump- ferret was noted in a dog town in Wind tion of 1080-poisoned prairie dogs by Cave National Park. Restocking of M. black-footed ferrets can cause fatal sec- nigripes in the National Park, and in ondary poisoning. other sanctuaries where prairie dogs Henderson et al (1969:31-37) re- thrive, such as Custer State Park and cently summarized the distributional Devils Tower National Monument, could status of M. nigripes in South Dakota. insure the survival of this interesting His census was based on questionnaires mammal. gi\'en to land owners, on a survey of Little is known concerning the repro- published literature and unpublished duction of the black-footed ferret but in U. S. records such as those located the young evidently are born in spring and Biological Survey files, and on examina- first emerge above ground early in July, tion of specimens housed in various mu- at which time family groups are most seums. Although the species was not un- frequently observed in western South common on the prairie along the eastern Dakota. of the Hills, a few records periphery only Specimens examined (4).—SOUTH DA- Hills are directly applicable to the Black KOTA: Pennington County: Spring Creek, proper. south of Rapid City, 2 (MHM). WYOMING: Weston County: Newcastle, In 1903-1904, P. Campbell ob- James 2 (USNM). a ferret in a town in served prairie dog Additional records.— (see text and Hender- Gillette Canvon, northeast of Elk Moun- son et a/.: 31-37, Appendix H). tain, Custer County. Another individual Mustela vison letifera Hollister was observed running along a road lo- cated in T. 2 S, R. 5 E, 5800-6000 feet, Mink Custer Clark in July County, by James Mustela vison letifera Hollister, 1913, Proc. the foot- 1965; this site may be within U. S. Nat. Mus., 44:475, 18 April (type hill region of Custer State Park, where locality, Elk River, Sherburne Co., Minne- habitat would be suitable for ferrets. sota). Troy C. Beach observed a M. nigripes Mink live along streams, edges of and lakes in all but the in a prairie dog town in Wind Cave ponds, marshes, National Park in the spring of 1922. southwestern part of the United States. in collections due to its valu- Upon completion of filming "The Van- Uncommon able Mustela vison is abun- ishing Prairie," Walt Disney Productions, pelt, fairly dant water courses the Inc., released two males and one female along throughout in the National Park on 30 December Black Hills. For example, "numerous near 1953 (Cahalane, 19.54:424; Garst, 1954: mink" are trapped Sturgis annually Dakota Conservation 23: 594). These were three of five ferrets (South Digest, (an adult and four young) trapped in a 12, March 1956). 132 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Grinnell (1875:80) saw no living under that name. Because of the great specimens while in the Hills, but ob- individual variation in the species and of served signs of mink along streams and because an adequate number speci- saw a skin that the owner said had been mens for analysis is lacking, I tentatively follow Hall in the Black taken "in the Black Hills." Bailey (1888: ( 1936) assigning 433) noted the presence of this species Hills specimens to Taxidea taxus taxus. in the Hills in November 1887. The badger is rather uncommon in A lactating female was taken along the Black Hills proper. Cahalane (1948: the edge of a beaver pond south of 249; 1951:209) reported that a pair was Tinton on 16 July 1961 by W. C. Stanley; released about 1940 in Wind Cave Na- to my knowledge, no additional data are tional Park, where tliriving prairie dog available for this species from the Black towns offer an ample food supply, and Hills, save for the specimens listed below. that this pair produced at least one litter Specimens examined (4).—SOUTH DA- of young. Koford (1958:31) noted that KOTA: Lawrence 2 nii S Tinton, County: several prairie dog tunnel systems in 6100 ft, 1. Pennington County: Rapid City, 1; Shirttail Canyon had been excavated by Martindale Ranch, 0.5 mi E Rapid City, 1 in 1955. In the of (SDMT); Spring Creek, south of Rapid City, badgers early morning 1 (MHM). 9 July 1968, I observed an adult female and two pups digging out a Cynomys Taxidea taxus taxus (Schreber) burrow on Bison Flats, in the National Badger Park; a Ranger-NaturaHst at the Park observed a female badger carrying a Ursus taxus Schreber, 1778, Die Saugthiere . . ., in her that description on pp. 520-521 (type locality, young pup mouth near prairie Labrador and Hudson Bay, restricted south- dog town on 20 May 1957 (Wind Cave west of Hudson by Miller and Kellogg, Bay National Park files). An adult T. taxus Bull. U. S. Nat. Mus., 205:747, 3 March was observed along the road 6 mi NW 1955). 5 1969 a road- Spearfish on August eating , Taxidea taxus—Rhoads, 1894, Amer. Nat. 28: killed Sciurus 524, June. niger. claims were in 1962 on Taxidea taxus is a relatively common Bounty paid inhabitant of the Northern Great Plains 37 badgers taken in the counties that the South Dakotan of surrounding the Black Hills, where it comprise portion the Black Hills from South Da- occurs throughout the Red Valley "race- (records kota Fish and Parks . One track," Wind Cave National Park, and Dept. Game, ) indi\'idual in the winter of 1964- other areas where fringes of prairie im- trapped 65 southwest of Hill was taken at an pinge upon the foothills. Being a rapid City elevation in excess of 6000 feet. and powerful digger, the badger pro- vides reliable evidence of its presence Specimens examined (6).—SOUTH DA- Meade 5.6 mi N 1 by its characteristic excavations and den KOTA: County: Piedmont, (SDMT). Pennington County: near Rapid sites, usually located on gently sloping City, 1 (FMNH); Spring Creek, south of Rapid hillsides. City, 1 (MHM); 4 mi S, 14 mi W Hill City, The genus Taxidea, with but one 1. Custer County: Dewey, 2 (USNM). Recent species, is distributed in centi'al hudsonica and western North America. Some years Mephitis mephitis Richardson ago, Viola S. Schantz began a revision, naming 10 new subspecies over a period Striped Skunk of several years, but a final report was Mephitis americana var. hudsonica Richardson, not issued; Charles A. Long currently is 1829, Fauna Boreali-Americana, 1:55 (type analyzing geographical variation of Taxi- locality, plains of Saskatchewan, Canada). — Univ. dea taxus. Schantz (1946:81) described Mephitis mephitis Jiudsonica Hall, 1934, California Publ. Zool., 40:368, 5 November. T. t. dacotensis from Folsom, Custer Co., South Dakota, just east of the Black Hills, The shiped skunk occurs throughout and included specimens from the Hills the United States. In the Black Hills, it TURNER: MAMMALS OF THE BLACK HILLS 133

mi ranges from the lo\^or elevations of the W Minnekahta, 3800 ft, 1; 5.5 mi E Minne- kahta, 4000 ft, 2. sunoimding plains, through the foothill WYOMING: Crook County: 3.5 mi NW ti-ansition zone, up to altitudes of about Sundance, 5000 ft, 1. 6200 feet. The species is at home in a Additional record—SOUTH DAKOTA: Fall variety' of habitats, but prefers forest River County: Minnekahta (USES card files). borders, wooded ravines, rocky outcrops, and brushy agricultural fields with asso- Family FE LI DAE—Cats ciated fencerows. It is no to sti'anger Of the three felids that occur in the the habitations of man, and although Black Hills, bobcats are by far the most in museum collections represented by common; mountain lions are less fre- only a few specimens, the species is com- quent, and lynx are but rare visitors to mon throughout the Black Hills region. the region in recent years. Grinnell (1875:80) first noted the pres- ence of this mustelid in the Hills. Felis concolor hippolestes Merriam Striped skunks often were observed Mountain Lion along roadsides and in the vicinity of Felis hippolestes Merriam, 1897, Proc. Biol. Soc. campgrounds. Residents of Nemo, Law- Washington, 11:219, 5 July (type locality, Wind River Mountains, near head Big ence County, indicated that more than Wind River, Fremont Co., Wyoming). 70 individuals had been killed in the Felis concolor hippolestes—Nelson and Cold- in 1966-1967; an adult community^ dump man, 1929, Jour. Mamm., 10:347, 11 No- female and an immature female were vember. obtained there on 30 1967. June Bailey Previously, Felis concolor occurred an individual near (1888:433) captured through much of North America, but it in a baited with the car- Rapid City trap has been extirpated in many parts of its cass of a Most prairie dog. remaining range and now occurs primarily in re- taken in the Black Hills have specimens gions poorly populated by humans. The been either killed on or roads, represent mountain lion feeds principally on deer skeletal material found in caves and and occasionally upon livestock. Due to rockv crevices. its role as a large predator, this species A juvenile was captured on 8 August has been persecuted to the point that 1894 at Custer by W. W. Granger. even within its present range, it is now Another immature Mephitis that was ob- rare and should be protected. Felis con- viously sick was captured along a road color is native to the Black Hills, although northwest of Minnekahta on 16 June it is uncommon and inhabits the more 1967. This indi\'idual offered no resist- remote and rugged areas. ance; possibly it had been poisoned or Members of the Custer Expedition of was rabid. A female taken near Nemo 1874 saw but one "panther," near the on 30 June had five placental scars, two head of Castle Creek (O'Hara, 1929: in the right uterine horn, and three in 252). Grinnell (1875:79), who accom- the left. panied Custer, believed that these felids were quite numerous and saw indications Specimens examined (30).—SOUTH DA- of their on several occasions: KOTA: Lawrence County: Nemo, 4700 ft, 2. presence Pennington County: near Rapid City, 8 "I . . . once found the partially de- (FMNH); Moon, 22 mi W Hill City, 6200 ft, voured remains of a deer thad had just 1; Ditch Creek Valley, 14 mi W Hill City, 6400 been left by one of these animals." Mem- Hill 4975 3 ft, 1; City, ft, 2; Spring Creek, bers of the Newton-Jenney Survey ob- mi E Hill 1 City, (UMMZ); Spring Creek, served two or three mountain lions but 1 Elk south of Rapid City, ( MHM ) ; 20 mi N none was killed (Dodge 1876:133). A Mountain, 1 (USNM). Custer County: Custer, female mountain lion and two young 3 (1 USNM, 2 AMNH); south of Otis, 1 were shot in November after (UMMZ); Wind Cave National Park, 1 1887, just the adult killed a mule deer (WCNP); Elk Mountain, 1 (USNM); Dewey, had large 3 (USNM). Fall River County: 1.75 mi N, 4.5 buck (Bailey, 1888:431) and Merritt 134 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Gary obtained a female near the head of different occasions in 1965, and on 2 Bear Creek in the Bear Lodge Moun- April of that year, fresh lion tracks were tains in 1890. identified in soft mud next to the pump Earl Bedell shot a lion near the head house. A lion also was observed about of Stockade Beaver Creek, Weston Co., this time on the ranch of Mr. Shirlev Wyoming, in 1930 (Mann, 1959:3) and McClure, southwest of Hot Springs. a female F. concolor was killed early in Specimen examined (1).—WYOMING: December 1931 near the bor- Wyoming Crook County: head Bear Creek, Bear Lodge der (Yomig, 1946b: 34). The latter indi- Mountains, 1 (USNM). vidual was known in the region for the and was believed to previous two years Lynx canadensis canadensis Kerr have raised at least one litter in this Lynx period. In a report written to the U. S. Biological Survey in 1932, Louis Knowles Lynx canadensis Kerr, 1792, The animal king-

dom . . 1:L57 restricted indicated that these were the only moun- ., (type locality to Pro\ince of Quebec, Canada, by Miller tain lions taken in the Hills in the pre- and Kellogg, Bull. U. S. Nat. Mus., 205: vious 25 years. 777, 3 March 1955). The most recent specimen taken in Lynx canadensis is typical of the the Black Hills to my knowledge was forested boreal of North shot by Ted Mann on 16 December 1958 heavily regions America, but formerly occurred spar- (Mann, 1959:5). He located fresh tracks in suitable habitat in the Northern in the snow on Elk Mountain, in Custer ingly Creat Plains region. Grinnell (1875:79) County. In the company of two neigh- and Dodge (1876:12.3) both indicated bors and three dogs, Mann set out in that this species previously inhabited the pursuit of the lion, which had killed a Black Hills, and there have been several doe and was eating on the hindquarters recent reports of lynx in the area. In when the dogs jumped it. The lion, a 1944, a Ivnx was killed in Meade Countv male that weighed 140 pounds and meas- and two more were taken in Pennington ured about seven feet in length, finally Countv (South Dakota Conservation Di- was treed and shot along Upper Dugout 12:15, A. H. Richard- Creek, about two miles south of the gest, May 1945). wrote Steam's Ranch. son, State Game Biologist, (pers. com.) that in the last 10 years he has In 1965, Fred A. Fichtner, District personal!)' examined two specimens shot from Newcastle, re- Ranger Wyoming, in the northern and western Black Hills. ported that there had been several re- I know of but three other individuals cent reports by ranchers of mountain in South Dakota. A was lion tracks between Custer and Elk captured lynx taken in Woods the Mountain, South Dakota; "Plaster casts Granger's along Missouri River, above Sioux Citv, in were made of one set, and positively 1875 A identified" January (Brackett, 1881:413). (letter to J. Knox Jones, Jr.). male (USNM 246.548) was obtained by Ranger A. W. Jones, Spearfish District, J. N. Martin at Bullhead, Corson County, noted the recent presence (1968) of a on 6 October 1925, and another was shot mountain lion in the vicinity of Big Crow by Mr. Leroy Johnson near Marindahl, Peak (pors. com.). Park Ranger Fred Yankton County, on 6 May 1962 (Lee, Devenport (pers. com.) indicated that 1962:21). Lynx canadensis presently in- two F. concolor were observed just in- habits eastern Montana (Hoffman and side of the west boundaiy of Wind Cave Pattic, 1968:.53), and has been taken National Park on 10 April 1964 by Dr. in the Laramie Ralph Hubbard and an Indian com- south of the Black Hills panion, of Wounded Knee, South Da- Mountains, Albany County, Wyoming kota. Ranger-Naturalists reported see- (Long, 1965:708) and in northern Ne- ing mountain lions in the Park on two braska (Jones, 1964:304). TURNER: MAMMALS OF THE BLACK HILLS 135

Lynx rufus pallescens Merriam Gulch. He placed the carcass on the Bobcat bank, set the trap, and captured three bobcats within a week (J. A. King, field Lynx fascidtiLs pallescens Merriam, 1899, N. notes). Stebler (1939:389) obtained a Amer. Fauna, 16:104, 28 October (type lo- of L. rufus Grace Cool- cality, S side Mt. Adams, near Trout Lake, specimen along in Custer State Park in Au- . Creek Skamania Co., Wasliington ) idge [Lynx rufa] pallescens—Elliott, 1901, Field gust 1934. The stream was bordered Columb. Publ. Zool. Mus., 45, ser., 2:297, with bur oak, quaking aspen, paper 6 March. birch, and other deciduous trees and The bobcat is widely distributed in shrubs. the Black Hills, being common in the Both of the females taken in Wind foothills, in forested canyons, along Cave National Park in early June 1924 in areas of wooded streams, and exposed were lactating. On 17 August 1967, I in the rimrock. It remains abundant shot a large female and two three-quarter Hills region in spite of predator control grown kittens at dusk as they were leav- 1311 bobcats measures. Bounties on ing a den at the base of a limestone out- those counties that com- were paid by crop just southeast of the National Park. prise the South Dakotan portion of the Thus, females of this species probably to 1968 Black Hills from 1962 (records give birth to young in late March or Fish from South Dakota Dept. Game, early April in the Black Hills. and Parks). In 30 years as a Federal Specimens examined (38).—SOUTH DA- Archie Howe of Custer killed Trapper, KOTA: Meade County: 6 mi E Piedmont, 1 in the Hills more than 450 bobcats (SDSU). Pennington County: near Rapid City, 1 south of (Popowski, 1952:6). ( FMNH ) ; Spring Creek, Rapid City, 1 (MHM); Palmer Gulch, SE Hill City, 3 Although he saw no specimens, Grin- (UMMZ); 4 mi S, 14 mi W Hill City, 1; un- nell indicated that this felid (1875:79) specified locality, 1 (NRW). Custer County: was common in the Hills region. Bailey S Otis, 1 (UMMZ); Custer, 3 (USNM); French Bowman (1888:432) noted the presence of L. Creek, near Custer, 5 (USNM); near Custer, 1 (USNM); Beaver Creek, rufus near Deadwood in October 1887, Ridge, south of Custer, 1 (USNM); Mayo, south of and collected one individual near Custer Custer, 1 (USNM); Streeter Ranch, 1.5 mi S, in 1888. July 5 mi E Wind Cave, 3550 ft, 3; Elk Mountain, 6 Evidently, the bobcat is an efficient 1 (USNM); Dewey, (USNM). WYOMING: Crook County: Otto, 1 predator on large game mammals. Young (AMNH); Leggot Canyon, Sundance, 1 that "A buck (1958:2.3) reported large (USNM); Person Canyon, Sundance, 1 antelope was no match for a bobcat (USNM); Sand Creek Canyon, 1 (USNM). which killed it one winter day of early Weston County: Newcastle, 1 (USNM). Black 1930 on the Wind Cave Game Preserve UNSPECIFIED LOCALITY: Hills, 3 (2 USNM, 1 WCNP). [Wind Cave National Park] in South . — Additional records ( see text also ) SOUTH Dakota." On 5 April 1954, George DAKOTA: Custer County: Bull Springs, 14 Barnes shot a 27-pound male that had mi NW Custer (Lamster, 1943:1). County Calumet Draw (South Dakota just killed a yearling white-tailed buck unspecified: Conservation Digest, 17:5, June 1950). near Wabash Spring, 5 mi N of Custer small also {op. c?Y.:84). However, game ORDER ARTIODACTYLA— is included in the diet of this species. Even-toed Stomachs of two females trapped in Ungulates Wind Cave National Park by Archie Artiodactyls in the Black Hills belong Howe in early June 1924 contained only to the suborder Ruminantia, and are rep- the remains of rabbits (U. S. Biological resented by three families with five gen-

files . six seventh Survey ) era and species. A species, introduced from In January 1946, a farmer found a the mountain goat, was additional dead beaver, with a trap attached, float- western montane areas. An the fallow has been intro- ing under the ice on a stream in Palmer species, deer, 136 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY duced from the Old World, but is not lowing account that partially indicated well established in the Hills as yet. Four the prior abundance of wapiti in the of the native species, wapiti, pronghorn, Hills: "On Elkhorn Prairie [Reynolds bison, and mountain sheep, previously Prairie] we came upon a collection of were extirpated or drastically reduced in liorns gathered together by the Indians. numbers, and subsequently have been Three lodge-poles had been set up in reintroduced from other areas. Artio- the ground so as to form a tripod, and dactyls comprise a varied assemblage of supported by these were a pile of [wa- hooved and horned, or antlered, mam- piti] horns 8 to 10 feet high. The horns mals that were economically important had been shed, and had apparently been to early explorers and pioneers. The pres- collected from the surrounding prairie ent-day game species still provide rec- [isolated prairie in the middle of the reation for outdoorsmen and much eco- Black Hills] and heaped up here by the logical information in the following Indians." Dodge (1876:132), describing accounts is drawn from conservation and a hunt along the northern rim of the game management literature. Black Hills, noted that a hunter "fol- lowed a large elk trail into a little valley" Family CERVIDAE—Elk and Deer that was densely vegetated and that at the report of his gun, "the whole valley Cervids are mainly browsers. Antlers was alive with rushing animals; at least are present at least in males and are a hundred, crashing through the brush, shed annually. This family is represented disappeared up the mountain side." Cer- in the Black Hills by two genera and vus canadensis was present in the lime- three species. stone countrv of the Black Hills, Bear Cervus canadensis canadensis Erxleben Lodge Mountains, Sundance Mountains, and Stockard Beaver and Wapiti along Inyan- kara Creeks, Weston Countv, Wvoming, canadensis [Cervus elaphtis] Erxleben, 1777, in 1885 (Murie, 1951:42),' and Bailey Systema regni animalis . . ., p. 305 (type (1888:434) noted that some were "said locality, eastern Canada, probal:)ly the vi- to occur still" in the Black Hills in 1887. cinity of Montreal according to V. Bailey, Proc. Biol. Soc. 15 Washington, 48:187, Near the turn of the century, J. A. November 1935). Allen (1895a: 263) indicated that the The wapiti, or American elk formerly \\'apiti had been extinct in the Black occurred throughout most of temperate Hills for se\'eral years. Thus, the original North America; it has been reintroduced population of wapiti (C. c. canadensis) in many areas where it previously had was mostly exterminated from the Hills been extirpated by man. The subspecies by 1900, and initial restocking with C c. C. c. canadensis was native to the Black nelsoni from Wyoming commenced. Hills, but, once it became extinct, the About 65 were stocked at Custer State area was restocked with C. c. nelsoni Park in 1912, but these escaped into the from Wyoming. Cervus canadensis may surrounding Hills and were replaced by be conspecific with the earlier-named an additional 25 in 1913; 50 more of these red deer, Cervus elaphus, of Eurasia large cerxids were obtained from Mon- (McCullough, 1969), but I retain cana- tana in 1916. Wind Ca\e National Park densis pending definitive study of their initiated a wapiti herd with 21 animals relationships. from Jackson Hole and Yellowstone Na- The Custer Expedition of 1874 saw tional Park in 1914. Currently there are 1200 in the State and only a few elk, but several were killed about wapiti Park, along Boxelder Creek near Nemo on 11 450 in the National Park (Cahalane, The herds are controlled August (Winchell, 1875:5.3; O'Hara, 1948:253). by

1929 : 252 . Grinnell 1875 : excess animals out of the confines ) ( 83 ) , who ac- drixing companied General Custer, gave the fol- of the parks into the surrounding Black TURNER: MAMMALS OI-^ TIIK HLACK HILLS 137

Hills, or by issuing hunting permits each vines of foothills, and in the more remote year. and rugged sections, lioth the mule PresentK, there are two herds, total- deer and white-tailed deer were an im- ing about 1000 elk, ranging through the portant somce of food for early travelers Black Hills (Evans, 1966:9). A small and settlers of the region; accounts of "northern herd" traverses the limestone Hayden (1856:79; 1859:706), Grinnell region along the South Dakota-Wyoming (1875:84), Dodge (1876:131), Bailey border, and a "southern herd" in habits (1888:434), and J. A. Allen (1895a:283) the area adjacent to the National and indicated that Odocoileus hemionus was State parks (A. H. Richardson, pers. present in the Hills in considerable num- com.). In the 1966-67 hunting season, bers. However, market hunting and 198 wapiti were killed by hunters in other factors essentially eliminated deer in Custer State Park, 40 ( 19 bulls, 15 cows, from most of South Dakota the early and six calves) were taken from the 1900's (Seton, 1929b : .332 ); only 225 deer, "northern herd," and 40 (21 bulls, 13 mostly O. hemionus, were taken legally cows, and six calves) were shot from the in South Dakota in 1920 (op. cit. -.250). "southern herd" (Rapid City Journal, A "buck law" and other legislation passed p. 9, 15 January 1967). in the period between 1911 and 1925 to in- There is much variation in the antlers enabled the populations of deer crease in abundance once more 1940 of this species in the Black Hills, and by deformations are not uncommon; some (Merwin, 1971:33). Currently, over- antlers are much flattened near the ex- population and o\'er-utilization of avail- able are the in tremities, being somewhat palmate in range major problems the ever Black Hills nature. Wapiti retire to dense forests managing increasing deer herd. during daylight hours, descending onto the \'alley meadows in early evening. The number of deer in the Hills in Bugling of males can be heard in the 1969 was estimated at 103,846—23 per- autumn during mating season, and caKes cent of which \\'ere O. hemionus (Rich- are born in late May or early June. ardson, 1969b: 25; Richardson and Russel, Pellet counts on a Specimens examined (4).—SOUTH DA- 1970:3). group (based KOTA: Lawrence County: 2 mi S Tinton, 365-day accumulation) conducted along 6100 ft, 2. Custer Countif: 1 mi NW Wind 107 belt transects in 1970 indicated a 16 Wind Ca^e National 1 Cave, Park, (WCNP). percent decrease in the Black Hills deer WYOMING: Weston County: 0..5 mi E population over that of 1969. Northern Buckhorn, 6150 ft, 1. Hills herd decreased 23.0 percent; cen- Odocoileus hemionus hemionus tral Hills herd decreased 14.6 percent, (Rafinesque) and southern Hills herd increased 5.3 Mule Deer percent (Thompson and Hausle, 1971:8). Mule deer are nati\'e to the Black Hills, Cervus hemionus Rafinesque, 1817, Anier. but the species currently seems to be Monthly Mag., 1:436, October (type lo- declining in abundance, occurring only cality, mouth of the Big Sioux River, South in small the Dakota ) . pockets throughout region, in Odocoileus hemionus—Merriam, 1898, Proc. with greatest concentrations the south- Biol. Soc. Washington, 12:100, 30 April. ern portion of the Hills. The ratio of white-tailed deer to mule deer varies The mule deer, or black-tailed deer, from area to area. The herd in Wind inhabits broken country, open plains, and brush or woods of western North Cave National Park and that of Fall America. An estimated 10 million mule River County are composed almost en- deer inhabited the continent when Eu- tirely of mule deer (F. Devenport, pers. com. whereas in the western section of ropean man arrived (Merwin, 1971:3.3). ) , there are This species occurs throughout the Black Custer County, equal numbers Hills but is most abundant in the ra- of each species. However, mule deer 138 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY predominate on the "West River Prairies" cinity of Annie Creek, Lawrence County, immediately adjacent to the Black Hills, serves as the main winter range for mule with the white-tail being essentially a deer, whereas white-tailed deer over- rixer winter the dweller of wooded bottoms ( Bever, on Hepler Range of the north- 1957:4); for example, 74 percent of the ern foothills. Both species utilize the 12,910 deer taken by hunters in this area Aztec Hill, McVey Burn, Crow Peak, in 1968 were O. hemionus (South Da- and Piedmont winter ranges. The criti- kota Conservation Digest, 36:4, Decem- cal period for deer in the Black Hills is ber, 1969). from January to April, and many cases Projected kill of deer in the 3517 of death due to malnutrition in winter

the have recorded et : square miles of South Dakotan por- been ( Gastler at., 1951 tion of the Black Hills in 1968 was 4.02 356). The severe blizzard in the winter deer per square mile and was composed of 1948-1949 resulted in the deaths of 40 of 31 percent white-tail does and 45 per- deer per square mile in the McVey Burn cent white-tailed bucks, 10 percent mule region (Berner, 1953a :4). Mule deer does and 14 percent mule bucks (Rose are especially susceptible to starvation 1970, Tables 4-6). and large herds in Spearlish Canyon must be fed concentrate South Dakota State Game Biologist high protein each winter due to overuse of the winter- A. H. Richardson (pers. com.) suggests that one of the main reasons for the de- ing grounds (Harris, 1952a:7), 1953:14; crease of mule deer in the Hills is the Brady, 1955:13; Leopold, 1956:4). Such in restricted areas further preference of this species for an open, aggregations the A of rolling type of habitat; in the past 20 to complicate problems. survey browse in and 1969 indicated 40 years, ponderosa pine has enclosed April May that in 2633 many former environs of this type square miles of wintering in the (Bever, 1959:4). In addition, white-tails range Hills, 92 square miles were 208 seem to be the more aggressive of the over-utilized; square miles were and 2333 miles two species in the Black Hills, forcing properly utilized, square were under-utilized out the mule deer. Also, relative per- (Richardson, 1969: centage of mule deer killed in the Hills 3). Over-utilization occurred on the by hunters is greater than that of white- McVey Burn, Crow Peak, and Piedmont tailed deer. Differential response of the ranges, and in the area of mountain two species to harsh weather conditions mahogany west of Custer. suggests another possible explanation of Rut extends from early November to the problem. For example, from 29 April mid-December. Fawns, usually twins, to 4 May 1953, snow in excess of 44 are born in the Black Hills in May and inches fell in the northern Hills. Odo- June. Deer lice (Lepoptena depressa) coileiis vir^iniamis, in the higher por- parasitize deer of the Wind Ca\e Na- tion of the Hills, migrated 10 to 25 miles tional Park herd externally (Wind Ca^'e down to the lower slopes and foothills National Park files). during the winter for better browse and Specimens examined (24).—SOUTH DA- more weather conditions. Mule equitable KOTA: Lawrence County: 1 mi S, 4 mi W deer did not migrate as readily, but Spearfish, 1. Pennington Cotintt/: Rapid City, simply shifted to more favorable expo- 1 (USD); Beaver Creek Vallev, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 1; 3 mi N, 4 mi E sures in nearbv steep canyon drainages Hill City, 1. Custer Cotinti/: Pleasant Valley (Berner, 1953b: 10; Harris, 1953:16; Ranch, 6 mi SW Custer, 2'(AMNH); Camp- Hodgins, 1956:14). hell's Ranch, Elk Mountain, 4800 ft, 1 Redhird Elk Southwest slopes, from which the (USNM); Canvon, Mountain, 6000 ft, 1 (USNM); Elk Mountain, 5 (USNM). snows melt most rapidly, and tops of Fall River Countt/: 1 mi N, 4 mi E Edgeniont, ridges, from which the snows are readily 2600 ft, 1. blown w are areas on which the by inds, WYOMINC: Crook County: Bear Lodge deer concentrate in the winter. The vi- Mountains [not 9524 (=U. S. Biological Sur- TURNER: MAMMALS OF THE BLACK HILLS 139

skull feet as noted hv vey number) Long, dant than any other animal" in the Black 1965:713—7242 feet is the hij^hest ele\ ation Hills. In strong contrast to these reports in the Blaek Hills], 2 (USNM). Weston is that of who wrote County: 3 mi E Newcastle, 1; unspecified lo- Bailey (1888:434), cality, 7. that "the black-tail [mule deer] is very common through the Hills but the white- Odocoileus virginianus dacotensis tail seems to be absent. I can find no one Goldman and Kellogg who has ever seen this deer here." Evi- White-tailed Deer dently, Bailey was in error in his obsera- tions, or confined his queries to the foot- Odocoileus virf^inianus dacotensis Goldman and hill because notations W. W. Kellogg, 1940, Proc. Biol. Soc. Washington, region, by in 1894 53:82, 28 June (type locality. White Earth Granger agreed with the previous Mountrail North River, Co., Dakota). accounts (J. A. Allen, 1895a:263). Occurring throughout much of North Odocoileus virginianus dacotensis America, white-tailed deer characteristi- differs from O. v. ochroura of western cally inhabit woodlands, forest edges, Wyoming (Long, 1965:714) and O. v. and thickets along streams. An esti- tuocroiirus to the east (Kellogg, 1956: mated 40 million white-tailed deer in- 44) in average larger size, heavier den- habited the continent when European tition, and paler color. However, there man arrived (Merwin, 1971:33). On the seems to be extreme variation in size Great Plains, this species occupies decid- within the geographic range of O. v. dac- uous riparian communities, but seem- otensis. Evidently, white-tailed deer in ingly avoids extensive open areas. Odo- the Black Hills region also are quite coileus virginianus is numerous and variable and may represent a broad zone of widely distributed in the Black Hills, intergradation with the adjacent sub- preferring timbered areas with adequate species. Jones (1964:319) felt that the cover and water that support a good alleged differences between subspecies forb and browse understory. The main did not appear to be great. The enigma wintering ranges are in the foothills, remains due to a lack of specimens for and on the McVey Burn in Pennington examination. County. This species is extremely adapt- Grinnell (1875:83) indicated that the able to human populations and is quite differences in size between the white- common in agricultural areas. In 1969, tails in the Black Hills and those on the there were in excess of 70,000 white- prairie were so great that hunters of the tailed deer in the Black Hills proper region thought the Black Hills deer to be (Richardson, 1969b: 25). a different species than that to the east. Grinnell wrote that this (1875:83) Dodge ( 1876:130) reported: "a few days species was abundant in the vicinity of previous to the arrival of General Cook, Castle Creek and the Elkhorn Prairie I killed near Rapid Creek two enormous [Reynolds Prairie] about the head of bucks, each of which, after disembowel- Elk all Creek, and through the north- ing, and having head and legs cut off, eastern portion of the Black Hills. He weighed nearly one hundred and thirty noted that 100 deer, principally of this pounds. In the hunt on Boxelder [with species, were killed by the command of General Cook], one of the guests killed General Custer on 9 August 1874 cast of a buck just as old and just as fat as those Custer Peak. Of 16 does killed by Dodge killed by me on Rapid, but which, (1876:121) between 15 August and 10 dressed in the same way, weighed September, only two had given birth to scarcely forty pounds." Dodge indicated fawns that season. Approximately 1000 that both the large and small forms were deer were taken by the Newton-Jenney found together in the same herd, as well Survey along Castle, Rapid, and Boxelder as animals representing gradations in creeks (op. cit. :122), causing Dodge to size between the mentioned extremes. state that O. virginianus was "more abun- Grinnell wrote as follows to Seton 140 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

(1929b: 321): "In 1874 when with Cus- birch, hawthorn {Crataegus succuhnta), ter's expedition to the Black Hills, L. H. hop hornbeam, redroot, hazelnut (Corij- North and I killed in the Black Hills lus cornuta), Oregon grape, bearberry, three or four extraordinarily small deer." wolfberry, snowberry (Symplwricarpos Numerous authors went on record in canadensis), Juneberry {Amehncheir Forest and Stream, and Recreation maga- canadensis), serviceberiy (A. spicata), zines during the period 1895 to 1900 as chokecherry, and russet buffaloberry having seen small white-tailed deer in (SJiepJierdia canadensis). Forbs utilized the Black Hills. Measurements of a pair by the Black Hills deer include pea vine In- American vetch of diminutive white-tail antlers from (Lat])yriis ochroleucus) , yan Kara Mountain, found in 1890 by {Vicia americana), blue aster (Aster Robert Ansley (Seton, 1929b: 316) are laevis), meadow sweet, wild pea (Lu- followed by those of a similarly-sized j)inus argenteus), pussy toes (Antenna- in the Mountains ria pair found Bear Lodge microphyUa) , wintergreen (PyroJa by Vernon Bailey in 1913 (USNM sp.), alfalfa, red clover, wild rose, soap- 202513): distance from burr to tip of weed, and various mushrooms, grasses

dis- lichens . blue- antler, 8 inches, 10 inches; greatest and ( Usnea sp. ) Kentucky tance between any two corresponding grass comprised over 90 percent of the points, 10 inches, 8.5 inches; distance grasses consumed (Hill, 1946:49). from base of one antler to base of second Progulski and Duerre (1964:27) antler, 2 inches, 1.75 inches; distance found that deer in the Hills were most from tip of one antler to tip of other active between one and five hours after 8.5 inches, 4.75 inches. A few antler, sunset. They indicated (op. cit. -.31) small adult white-tails still are unusually that activity was most concentrated in killed by hunters each season in the lush meadows that were interspersed Black Hills (Over and Churchill, 1945: with brushy cover along streams. Rich- 51; Bever, 1957:4). ardson and DeMarce (1967:8) analyzed Nutrition may be a possible explana- the numbers and distribution of deer in tion of the problem. Both white-tailed the Black Hills in 1966 and 1967, based deer and mule deer in the Hills are on a 365-day accumulation of deer pel- smaller in body size than are prairie rep- lets. They found 19.7 percent of the deer resentatives of the same respective sub- occurred in the northeastern section, 47.7 species (Berner, 1953c: 12). Over-popu- percent in the northwestern-central sec- lation and over-utilized range are char- tion, 22.0 percent in the southeastern- acteristic of the Black Hills herd, but central section, and 10.6 percent in the not of the prairie herd. Food habits are southeastern section of the Hills. the same for mule deer and essentially Mr. L. Dunn of Sturgis shot a white- white-tailed deer in the Black Hills, and tailed buck with palmated antlers near have been Hill and Harris reported by Moskee, Crook Co., Wyoming in 1965; Hill Gastler et al (1943), (1946), there ^^'ere 13 points on a side and the (1951), Bever (1954), and Schneeweis antlers much resembled those of a small As (1968). determined by pellet counts, moose (Lee, 1965:19). In the autumn deer utilize mixed aspen-pine communi- of 1968, Dr. Calvin Schad shot a white- ties to a greater extent than pure stands tailed buck in the Black Hills that had of either or Kranz and Peter- pine aspen ( three antlers: a normal four-point antler sen, 1970:5). Important browse species on the left side of the skull, and two based on of browse ( observation, surveys three-point antlers, arising separately, and of stomach con- utilization, analysis on the right side (South Dakota Con- tents of 320 deer taken in the northern servation Digest, 36:13, April, 1969). section of the are as follow: Hills) Rut begins early in November in the ground juniper, crcx^ping juniper, (juak- Black Hills, reaching its maximum dur- ing aspen, bur oak, ponderosa pine, paper ing the latter part of the month. Does TURNER: MAMMALS OF THE BLACK HILLS 141 first breed as yearlings and usually bear and is represented in the Black Hills re- twin fawns from late May to early July. gion by one Recent species. Both sexes On one occasion I witnessed a doe with of this species possess permanent bony three fawns near Custer. Mule deer- outgrowths from the skull that are cov- white-tailed deer hybrids have been re- ered with highly keratinized horny ported from the Hills but none have sheaths; these sheaths are shed and re- been authenticated (Bever, 1957:5). placed annually. Pronghorn are grazers During late summer and early au- and inhabit the open flat country and tumn of 1952 and 1956, outbreaks of gently rolling upkmds of the Great epizootic hemorrhagic disease (EHD) Plains. killed hundreds of white-tailed deer and an occasional mule deer in western South Antilocapra americana americana (Ord) Dakota, including Meade, Pennington, Pronghorn Custer, and Fall River counties (Shope Antilope americana Ord, 1815, in Guthrie, A et al, 1960; Pirde and Layton, 1961; new geograpliical, historical, and commer- Trainer, 1964). Of the two viral strains cial . . grammar ., ed. 2, 2:292, 308 (type known to be causative agents, the "Black locality, unknown; noted in the original Hills strain" is far less virulent than the description as found "on the plains and high-lands of the Missouri"). "South Dakota strain" that causes epi- Antilocapra americana Ord, 1818, zootic outbreaks among prairie deer Jour. Phys. Chim. Hist. Nat. et Arts, 87:149. (Parikh, 1970:1). These EHD-causing strains currently are under investigation Pronghorns now are reduced in num- by virologists at South Dakota State Uni- bers over much of their former range. versity. Deaths by automobiles is another An estimated 30 to 40 million once mortality factor. For example, more than roamed over the western half of the 30 deer were killed on highways in the United States (Long, 1965:716). Agri- northern Hills from 16 September to 12 cultural pursuits of man and settlement October 1952 (Harris, 1952c: 16). of former pronghorn range contributed to a decline in the abundance of this Specimens examined (29).—SOUTH DA- KOTA: Lawrence County: 3 mi S, 3.5 mi W species (Nelson, 1925:3). Hoffman 1. 16 mi Spearfish, Pennington County: W (1877:100) indicated that a fatal epi- HiU 6600 ft, 1; 5 mi W Hill 2; City, City, demic raged among pronghorns in the near Pactola, 1 (USD); 6 mi NE Keystone, 2; summer of 1873, and that J. A. Allen 4 mi N, 2 mi E Keystone, 4800 ft, I; 20 mi N Elk Mountain, 1 (USNM). Custer County: estimated that from three-fourths to nine- Pine Creek, 1 mi N Harney Peak, 1 (SDSU); tenths of the population may have been Elk 2 Mountain, (USNM). destroyed in some areas. Coincident WYOMING: Crook Bear County: Lodge with this epidemic, an epizootic disease Mountains, 13 (USNM); Lost Canyon, 5000 killed most of the government stock and ft, 1 (UW); unspecified locality, 2 (AMNH). Indian in western South Dakota. Weston County: vmspecified locality, 1. ponies Additional records.—SOUTU DAKOTA: Hoffman {loc. cit.) noted that "If the Lawrence County: Elk Creek (Harris, 1952b: horse epidemic was not the cause of Custer Bull 14 mi 12). County: Springs, NW fatality among the antelope, it is at least Custer WYOMING: (Lamster, 1943:14). in very remarkable coincidence." Nearly Crook County: Moskee (Lee, 1965:19). 64 percent of the Indians at this time Crook's Tower ( Chicago Tribune, 12 Novem- ber 1941). UNSPECIFIED LOCALITY: were affected with cerebro-spinal menin- Black Hills (Hoffman, 1877:100). gitis, and suffered a 10 to 12 percent mortality. The Indians procured their Family ANTILOCAPRIDAE— drinking water from the same pools Pronghorn where their ponies drank. The pronghorn is not an antelope but On 16 October 1804, Indians told is a member of a unique family that has Meriwether Lewis and William Clark evolved exclusively in North America that the pronghorn were then on their 142 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY fall migration west to the "Black Moun- Present rangeland management in

winter 1927 : western South is tains" to spend the ( Bailey, Dakota oriented toward 28). Hayden (1862:150) wrote: "In the grasses and other graminoid plants. Be- beginning of winter they [pronghorn] cause native woody range plants and may be seen for days following each forbs are high in fats, oils, carotenes, other in files (if not disturbed) on their protein, and other constituents necessary way towards the Northwest, leaving the for winter maintenance of both big game prairie for the more rugged portions of and domestic livestock. West (1971:4) the country near the Black Hills or the suggested the need for re-establishment foot of the mountains. In the spring, of these browse-plants species. The cur- usually about March, they may be seen rent population of pronghorn on that returning again, and distributing them- portion of the prairie adjacent to the selves over the open prairie." Seton Black Hills is about 82 percent of esti-

also that mated . (1929b: 421) reported prong- carrying capacity ( see Table 28 ) horn of the open country flock to the In managing pronghorn, the Depart- Black Hills "from all points on the com- ment of Game, Fish and Parks divides pass" to winter. sections of South Dakota into "Hunting In his review of the status of the Units" that can be censused and adminis- pronghorn. Nelson (1925:3) recorded 11 tered independently. Units that surround bands, totaling an estimated 680 indi- the Black Hills are as follow: Unit 3, viduals, as being resident in South Da- western Butte County (just north of the kota, and Berner (1954:2-3, 1955:2-3) Hills); Unit 4, eastern Butte and north- summarized the history of the prong- western Meade counties; Unit 9, southern horn in South Dakota. At present, spring Meade, central Pennington, and western inventory of pronghorn is carried out by Custer counties; Unit 14, eastern Fall the South Dakota Department of Game, River County; Unit 6, western Fall River Fish and Parks each year. The aerial and southwestern Custer counties. Table count, conducted from 3 June to 8 July 28 summarizes the composition of the 1970, indicated that the population in 1970 pronghorn populations in these re- western South Dakota numbers about spective units (see West, 1971: Tables 25,000 individuals, representing a six per- land 2). cent increase over the 1969 estimate A herd of about 300 pronghorn cur- (West, 1971:1). A census of pronghorn rently is maintained in Wind Cave Na- in counties east of the Missouri has not tional Park. Dr. W. T. Hornaday of the been attempted for several years, but Bronx Zoo, and the Boone and Crockett 302 individuals were killed by hunters in Club, contributed the monev for the these areas in 1968 (West, 1970:2). initial restocking of the Park in 1914.

Table 28.—Estimated composition of the 1970 populations of Antilocapia

americana in Hunting Units that border the Black Hills (West, 1971 ).

•« to

Hunting Unit TURNER: MAMMALS OF THE BLACK HILLS 143

With permission of the Canadian authori- Numerous pronghorn were captured in ties, 13 six-month-old kids were pur- northwestern South Dakota for release chased from Mr. Blazier of Brooks, Al- on the Lower Brule Indian Reservation, berta, for $125.00 eaeh. These were Lyman County, and on the Igloo Black reared for a short time, then were liber- Hills Ordnance Depot, Fall River ated in the Park that October (Nelson, County, in 1963 (South Dakota (Conser- 1925:9; Grinncll, 1929:139; Wind Cave vation Highlights, 1964). One hundred National Park files). Disease and preda- and twenty pronghorn were live-trapped tors reduced the number to eight by in Wind Cave National Park in 1963 for 1915, and nine more were purchased release on the Lower Brule Indian Res- from Alberta in 1916. Coyotes and bob- ervation, on private lands in Gregory cats continued to take a high toll of the County, and for the Kansas Game and pronghorn until predator)' animal hunt- Fish Commission for release in southern

the Kansas National Park files . ers were detached to the Park by ( Wind Cave ) U. S. Biological Survey; these hunters These are but a few instances of artificial eliminated sexeral hundred predators mixing of the herds in South Dakota. (Nelson, 1925:17). By 1924, only six On 2 September 1964, the "Head- animals all does but a ( ) survived, young quarters Fire" burned approximately buck was captured near Bisner, Nevada, 4000 acres of Wind Cave National Park. in The and released in the Park July. The kid crop of the next year was much 1929 herd numbered only 28 by (Grin- reduced from that of previous years. ncll, 1929:140), but 40 individuals sur- Pronghorn moved into the burned area the vived the disease and drought of and fed mainly upon pricklypear cactus 1930's. were well- Pronghorn finally just prior to the peak of the reproductive established in the Park by 1948 and num- season. False gromwell (Litlwspermum the bered about 150 animals, in spite of multiflomm) is known to inhibit ovula- the loss of 35 individuals to poachers in tion in some grazing mammals, and previous year (Cahalane, 1948:256). It Ranger-Naturahsts at the Park speculate control of carnivores in is probable that on the possibility that cacti, from which is of merit the region no longer owing the spines have been burned, might have to the natural balance that now exists had a similar eff^ect on the reproductive between and from various re- predator prey. potential of pronghorn ( Management programs have involved ports on file at Wind Cave National in transporting live-trapped individuals to Park). Breeding takes place Septem- areas of low population density for re- ber and young, usually twins, are born lease. These introductions have been in late May and June. numerous and difficult to trace. Twelve Specimens examined (10).—WYOMING: kids introduced into Custer State were Crook County: Sundance, 7 (USNM). Weston from Indian Creek Butte Park Flats, County: 3 mi S, 1 mi E Newcastle, 1; imspeci- 10 were added County, in 1916, and more fied locality, 2. Conservation Di- in 1919 ( South Dakota gest, 36:3, 1969). Eight pronghorns Family BOVIDAE—Bison, Sheep, in Meade were released trapped County AND Goats in Custer State Park on 4 February 1952 (Berner, 1952:8). Fift\'-one pronghorn This family of large herbivores is were transplanted from Custer State represented in the Black Hills by thi-ee Grant Park to Leola Hills, near Milbank, species. Two of these formerly were na- County, in 1959 (South Dakota Conser- tive to the region and have been reintro- vation Highlights, 1960), and 126 were duced, one being replaced by a non- transferred from Butte Count}^ to Day native but similar subspecies. The third and Edwards counties in 1961 (South representative is a non-indigenous spe- Dakota Conservation Highlights, 1962). cies established by man. 144 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

Bison bison bison (Linnaeus) western South Dakota and adjoining of to white Bison portions Wyoming man, rather than to the Sioux Indians. If the ed. Bos bison Linnaeus, 1758, Systema naturae, Sioux often hunted in the manner de- 1:72 (type locality, "Quivira," central 10, scribed then their contribu- Kansas—see Hershkovitz. Proc. Biol. Soc. by Warren, decimation of bison in Washington, 70:32, 28 June 1957). tion toward the the Black Hills was indeed. B[ison'\. bison—Jordan, 1888, Manual of the region great

vertebrate animals . . ed. 337. ., 5, p. General W. F. Raynolds (1865:27), The number of bison that once in the course of leading an expedition to roamed the plains of North America has explore the Yellowstone in 1859, caught been estimated between 60 and 100 mil- first sight of the "lords of the prairie" Hon by various authors. The bison evi- as his command approached the Black dently was common on the broad, un- Hills. When military posts just to the Hills dulating prairies surrounding the Black west of the Black were abandoned Hills in the early 1800's, but at the time in 1868, bison were still abundant, 50 of the first exploration of the region by tons being killed for garrison use in one European man, it was rare to see even day; yet, by 1871 no bison survived in an occasional wanderer. Bison had eastern Wyoming (J. A. Allen, 1877:544).

east of Missouri Grinnell 1875 : 84 while wholly disappeared the ( ) , accompanying Biver prior to 1870, but great herds General Custer in 1874, observed no live roamed the Coteau des Prairies west of bison in the Black Hills, but found a few the James River in South Dakota as late skulls with the hide still attached, indi- of this bovid. as 1866 (J. A. Allen, 1877:538). This cating recent presence species was decimated in the region be- Much later, Fryxell (1926:103) also dis- tween the Grand and Cheyenne rivers covered bison remains in the Black Hills. about 1869, although occasional strag- Bison in the Black Hills today are the glers frequented the plains toward the the result of reintroduction. In 1913, Black Hills until somewhat later (Hoff- National Bison Society provided $26,000 man, 1877:101). with which to purchase bison to restock the Wind Cave National Game Reserve. When Lt. G. K. Warren (1859:630) and his command traversed the western W. T. Hornaday secured the nucleus herd of 14 animals bulls and edge of the Black Hills in September (seven seven cows from the Bronx Zoo 1857, they encountered a large force of ) through the New York on 28 Hunkpapa and Miniconjou Sioux in the Zoological Society November 1913. These bison were vicinity of Inyan Kara Mountain, Crook orig- the zoo from WilHam County, Wyoming. Warren gave the inally acquired by C. of the October Mountain following account: "In the first place, Whitney Game Preserve in the Berkshire Hills they [the Indians] were encamped near near in 1903. large herds of buffalo, whose hair not Lenox, Massachusetts, in them from being sufficiently grown to make robes, Whitney, turn, purchased H. K. of Moosehcad Ranch at the Indians were, it may be said, actually Gliddon, one of the bulls had herding the animals. The intention of Jackson, Wyoming; the Indians was to retain the buffalo in been captured wild in the panhandle of Texas Colonel Charles "Buffalo" their neighborhood till their skins would by J. bulls and four cows from answer for robes, then to kill the animals Jones. Two Yellowstone National Park were added by surrounding one band at a time and to the herd in 1916. completely destroying each member of it." Fearing that Warren's expedition The Custer State Park bison herd, would disperse the bison, the Indians now the second largest in the United forced it to turn back southeastward. States, was started in 1914. The South Most writers have attributed the rapid Dakota State Legislature appropriated extermination of the bison over south- $15,000 to stock the reserve with 36 ani- TURNER: MAMMALS OF THE BLACK HILLS 145 mals (Six bulls, 18 cows, and 12 calves) KOTA: Custer County: "Wind Cave Game 1 Vail River Hot that were purchased from the James Reserve," (USNM). County: Springs, 1 (USNM). estate, Fort Pierre, South 'Scotty" Philip Additional records (see text also).—SOUTH 'Dakota. This herd originated from five DAKOTA: Custer County: French Creek cahes captured along the Grand River (Grinncll, 1875:84). near Fort Bennet, in northwestern South Oreamnos americanus missoulae Dakota, by Peter Dupree in 1882. The A. Allen Dupree herd increased to 60 by 1900, J. when Philip purchased the entire stock. Mountain Goat An additional 60 bison were obtained by Oreamnos montanus m,issoidae J. A. Allen, 1904, the State Park from the Pine In- Ridge Bull. Amer. Mus. Nat. Hist", 20:20, 10 Feb- South Da- dian Reser\ation, Rosebud, ruary (type locality, Missoula, Missoula Co., ikota, in 1951 (South Dakota Conserva- Montana). — tion Digest, 20:23, December, 1953). Oreaninos americanus missoulae Hollister, Proc. Biol. Soc. Washington, 25:186, Presently, approximately 350 animals 1912, I 24 December. 'range over 27,220 acres in Wind Cave National Park, and about 1550 animals Mountain goats are not native to the range over 72,000 acres in Custer State Black Hills (Swift, 1941:441) and other- Park. These numbers are maintained by wise have not become established east culling and selective harvesting. For ex- of the Continental Divide. In February ample, 276 bison were auctioned by Cus- 1924, U. S. Senator Peter Norbeck and ter State Park in 1968 at a live sale and State Game Warden L. C. Hawley ob- 138 others were butchered Da- a (South tained six goats, four females ( yearling kota Conservation Digest, 36:29, Decem- and three adults) and two males (a ber 1969). Each autumn, usually in yearling and an adult), from near Banff, October, the bison in Wind Cave Na- Alberta, Canada (Gilhland, 1968a :3). tional Park are rounded up by use of These were placed in a 20-acre pen in helicopters, horses, and vehicles, herded Custer State Park, but an adult female into a corral, branded, and treated for and a yearling male escaped the first ani- Bangs disease (Brucellosis). Excess night. Warden Hawley indicated to mals are removed for slaughter and sale Hanson (1950:21) that no young were in cooperation with the State of South born to the captive goats, but Harmon Di- South Dakota Conservation : stated that the Dakota ( ( 1944 149 ) remaining gest, 20:3-5, 7; Griffith, 1958:16-19). In four increased to eight by 1929, at which 1968, 10 hunting permits ($500 each) time all escaped (when a tree fell across were issued to hunt bison in Custer State the pen fence) to the rugged Elkhorn Park. country near Harney Peak. The herd and the Throughout most of the year, bulls increased to 300 by 1949, popu- lation has remained at about that level remain solitary, or in small groups. About to the time. Goats were trans- late July, the bulls begin to join the present cow-calf herd and mating occurs, usu- planted to Spearfish Canyon, north of several but none has ally in August. Calves are born between Savoy, years ago, since then mid-April and June, after a gestation been observed there (South period of about nine months. Presently, Dakota Conservation Digest, 22:11, July, the Wind Cave herd is being studied 1955). extensively by Milo Schulte, Iowa State Oreamnos americanus is stenoecious University, Ames, Iowa. Garretson in the Black Hills, inhabiting only the (1938) and Roe (1951) have written Mount Rushmore-Needles-Harney Peak valuable accounts concerning the past area in the southern part of the crystal- history, disti'ibution, and natural history line central basin, where moss- and of l)ison. lichen-co\ ered granitic ridges are thrust to the sur- Specimens examined (2).—SOUTH DA- upward nearly perpendicular 146 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY rounding forest. The range of this spe- Hills evidently establish individual home cies in the Hills comprises approximately ranges and remain within these limits 32,000 acres, of which only 2080 acres the year around (Hanson, 1950:74). Al- are "primary range" (Schara, 1967:22). though home ranges overlap one another Population density on the primary range to varying degrees, the goats seldom is about 12 goats per square mile, and gather into groups except during rut. approximates four goats per square mile Mating usually occurs in October and on less desirable range (Hanson, 1950: November. After a gestation period of 27). about 180 days, one (usually) or two are born in Harmon (1944:149-151) and Hanson (occasionally) young April on the food habits or May in a secluded place amid dense ( 1950:55-63) reported timber that is with rock- and natural history of the Black Hills intermingled Kids are mountain goat herd. Feeding takes place piles (Hanson, 1950:42). weaned the but at short, irregular intervals throughout following August, may the mother until the next the day, but most intensive foraging oc- accompany curs from late afternoon to dark. Food young is born. Reproduction does not occur until two of Of 233 plants generally include mosses, lichens, years age. goats from 1948 to grasses, bracken fern {Pteridhwi aqtiili- sampled Julv February Hanson 138 mim), yarrow (Aclnllea lamilosa) cur- 1949, (1950:28) reported 36 and 59 rant (Rihes inehrians), fleabane {Erig- adults, yearlings, young. Shed of the outercoat eron sp.), bearberry, serviceberry, choke- long begins in mid- the worn outer fur is com- berry, hazelnut, buffaloberry, meadow April; shed The shorter sweet, paper birch, willows, quaking pletely by early July. undercoat molts in the latter of this aspen, ponderosa pine, white spruce, and part A new outer coat is ground juniper. The most important period. fully grown 1950:36- food of all seasons, both in preference by mid-September (Hanson, and in quantity consumed, is the tree 38). lichen known as "old-man's beard" A recent survey by Arthur H. Rich- sites and food ardson State Biolo- ( Usnea harbata). Feeding (pers. com.). Game habits vary with season. In spring, the gist, indicated that the browse plants goats feed on succulent graminoid vege- have been severely overused, with pre- tation in small grassy parks, near rocky ferred food species being drastically re- areas, along the bottoms of canyons. In duced in the primary range of the goats. summer and autumn, they forage among Trapping operations indicated that the small coves of aspen and birch scattered general health of the herd was declining over slopes and ridges. In winter, the rapidly. In order to combat a feared goats retire to the timber, or descend to population crash, the first mountain goat protected areas such as the gulches of season was held in 1967. Forty-six goats Grizzly and Pine creeks, there competing were taken by hunters in the next two with deer and elk for available food. years, several of which were large Winter foods consist of 60 percent mosses enough to be registered with the Boone and lichens, 20 percent bearberry, 10 per- and Crockett Club. On 8 March 1956, cent pine twigs and needles, and 10 per- a large male scoring 53K points, was cent miscellaneous browse, grasses, and found dead along Grizzly Creek, 3 mi E weeds. Goats frequently visit salt blocks Harney Peak (South Dakota Conserva- placed atop Harney Peak by the U. S. tion Digest, 24:5, August, 1956). Forest Service. Visceral samples colected from 28 In the Black Hills, district seasonal goats in the 1967 and 1968 hunting sea- migrations are limited or nonexistent, sons revealed that the population was aside from short altitudinal movements heavily infested with endoparasites correlated with procurement of food (South Dakota Conser\ation Digest, (Bever, 1955:4). Mature goats in the 36:5, December, 1969). Three kinds of TURNER: MAMMALS OF THE BLACK HILLS 147 cestodes {Moniezia J)cneden\, Taenia species that now is extinct (Cowan, 1940: hijdatigena, and Thi/sanosoma aciinoides 542), was native to the Black Hills. In- in the intestines) and 11 kinds of nema- troduction (Jackson, 1944:28) of the todes (Pwtostronyhis nishi and P. stUesi morphologically similar \\{\st(>rn subspe- in the lungs; 0.stcrtaij,ia circuincincta, O. cies, O. c. canadensis, accounts for the ostertagL O. trifitrcata, TricJiostrogyhis present occurrence of moimtain sheep in axei, and T. coliuiibriformis in the abom- the Hills. Jwlvetianus and N. asa; Neniatodinis Honess and Frost (1942:4) indicated macidosus in the small intestine; Oesoph- that O. c. auduhoni may be indistinguish- agastonium vemdosum and TricJiuris sp. able from O. c. canadensis. Lacking identified from in the caeca) ha\'e been specimens, I follow Cowan (1940:542- these visceral and samples (Boddicker 543), who critically examined material Hugghins, 1969; Boddicker, 1970; Block, from throughout the range of the species, Ticks 1970- Boddicker et al, 1971). in applying the name auduhoni to the (Dermacentor andersoni) and chewing native mountain sheep of the Black Hills lice (Bovicola. oreatnnidis) parasitized and adjacent areas. However, Cowan mountain goats in the Black Hills ex- (1940:543) noted that "O. c. auduhoni et and ternally (Boddicker al, 1971:95), based as it is on slight cranial characters million of residues of 0.59 parts per presented by a small number of speci- chlorinated hydrocarbons ( heptachlor mens is to be regarded as a weak race." and DDE were dieldrin, DDD, ) epoxide, In 1833, Maximillian reported that isolated from fat of 13 kidney goats (op. the Manitarii Indians usually went to the d^:101). Black Hills and associated mountainous and Overcrowding of available range tracts to hunt mountain sheep, killing a heaxy lungworm infestations that result hundred or more in a season (Thwaites, in diseases of the lungworm-pneumonia 1906:246). However, the maps of Maxi- complex probably are major factors in millian labeled the Killdeer Mountains controlling population levels of mountain and the Badlands of the Little Missouri goats in the Black Hills. A treatise on River (both in North Dakota) as the cur- this species and its management "Black Hills" (Bailey, 1927:25). Mem- rently is in press (A. H. Richardson, The bers of the Newton-Jenney U. S. Geologi- Rocky Mountain Goat in the Black Hills. cal Survey shot a mountain sheep along South Dakota Dept. Game, Fish and a small tributary of Boxelder Creek on Parks, Bull. 2, Pierre). 20 July 1875 (Dodge, 1976:129). Seton (1929b: that mountain Specimens examined (2).—SOUTH DA- 535) reported KOTA: Cttster County: Harney Peak, 1 sheep were essentially gone from the (SDSU). Unspecified County: Black Hills, 1 Black Hills by 1887, although a few lin- (SDSU). gered on until 1899 when the last indi- vidual about which he had information Ovis canadensis auduboni Merriam was killed. W. S. Phillips, however, in- Mountain Sheep dicated that mountain sheep ranged in the summer of 1890 from Ovis canadensis auduhoni Merriam, 1901, Proc. Sundance, Wy- Biol. Soc. Washington, 14:31, 5 April (type oming, to the Inyan Kara and Bear Badlands locality, Upper Missouri, probalaly Lodge mountains, all along the western the and White rivers. betxveen Cheyenne slopes of the Black Hills (Grinnell, 1904: South Dakota . ) 1895a: re- 3.34-335), and J. A. Allen ( 263) Mountain sheep formerly inhabited ported a small herd in the vicinity of of the foothills and prairie breaks, as well as Harney Peak near the turn century. were seen Seton loc. cit. mountainous areas, in the western United No sheep by ( ) Hills in States, but now occupy only rough buttes when he xisited the the summer of about 200 O. and canyons in the more rugged moun- of 1902, but a band tains. Ovis canadensis auduhoni, a sub- canadensis still survived to the east in 148 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

the White River Badlands, near Porcu- (Nachtigal, 1961:4), and 22 from Colo- pine Hill in 1909 (Charles, 1964:23). rado were received at the Badlands Na- The idea that native sheep were initially tional iMonument in 1963 (Charles, 1961: prairie dwellers (Grinnell, 1928:1) and 5; Conservation Highlights, South Da- were driven to the mountains seems to kota Dcpt. Fish, Game and Parks, 1964). be more closely associated with audu- In January 1967, a single ewe with honi than with any of the other races one lamb was observed in the Red Valley (Buechner, 1960:21). in Wind Cave National Park. A skeleton was found in the A herd of eight Ovis canadensis, ob- same area in the sum- mer of tained from Alberta, Canada, in 1922 by 1968, creating some doubt as to U. S. Senator Peter Norbeck, was con- the survival of the ewe and lamb. On 15 nine mountain fined in Custer State Park, but a sore- January 1969, sheep ( one mouth virus all but eliminated them bv adult male, seven adult females, one ju- 1961. A new herd was brought in from venile male) were transplanted from Du- Montana about 1965 and now consists of bois, Wyoming, to the northwest side of the In\'an Kara from 50 to 70 sheep (A. H. Richardson, Mountain in sec. 24, T. 49 R. 63 Crook pers. com.). Some indi\'iduals, mainly N, W, County, Wyoming. rams, escaped into the surrounding Hills. Eight of these ascended the mountain but one female across the Possibly, additional individuals from slope, escaped As of 8 October the mature other herds introduced into nearby areas prairie. 1969, ram and one female four in western South Dakota may have died; howe\'er, new lambs have been observed (F. C. reached the Black Hills. Twelve moun- Windsor, pers. com.). tain sheep obtained from Alberta were Specimens examined (1).—SOUTH DA- released on the Claude Olson Ranch, KOTA: Custer Cotmtij: Wind Cave National Slim Buttes, Harding County, in 1961 Park, 1 (WCNP).

SPECIES OF UNVERIFIED OCCURRENCE

SPECIES INCORRECTLY REPORTED specimen of the \'agrant shrew in the FROM THE BLACK HILLS Museum of Natural History, The Uni- versity of Kansas, from a localitv 3 mi Sorex vagrans monticola Merriam, NW Sundance, 5900 ft, Crook Co., Wy- 1890.—Baker and Findley (1953:382) oming. There is no evidence that such a reported a specimen of the vagrant shrew specimen e\'er existed at the Museum; from 2.5 mi N Fairburn, Custer Co., however, there is an additional South Dakota. They wrote as follows: speci- men of Sorex cinereiis Jiaydeni beyond "We have tentatively referred this speci- the number indicated by Long for the men to S. t). monticola Merriam on geo- specified localit)^ Findley ( 1955 ) revised graphic grounds, although certain cranial the species Sorex vagrans and listed no measurements taken show the shrew to specimens from Crook County; yet. be much smaller than other shrews of Long (1965:520) wrote, ". . . my findings this subspecies. . . ." Subsequently these agree with his [Findley] concerning S. same authors (Findley and Baker, 1956: vagrans in Wyoming." Thus it appears 543) assigned the same specimen to that Long's report of the vagrant shrew Sorex nanus (see account of that spe- occurring in the Black Hills is in error. cies). Findley (1955:26) suggests that a segment of the ancestral S. vagrans Myotis evotis evotis (H. Allen, 1864.) stock have in the Black might persisted —The long-cared m\'otis has been re- Hills the during Sangamonian interval. ported from the Badkmds to the south- Sorex vagrans obscurus Merriam, east of the Black Hills and from Hard- 1891.—Long (1965:522) reported a ing County to the north. It also is known TURNER: MAMMALS ()i< HIE BLACK HILLS 149 from much of Wyoming, excluding the gust 1857 "along a wagon road to Bridger northeastern portion of the state. Pre- Pass" in the region now known as the vious reports of M. evotis from the Hills Laramie Mountains, Wyoming, and the were erroneous. Of three specimens heath(>r \'ole does not occur in the Black listed by Stebler (1939:389) from Custer Hills. State Park, South Dakota, one is Myotis A. tlujsanodes, and the other two are M. Zapus princeps princeps J. Allen, 1893. —When the list keenii. One reported from 1.5 mi E composing general Weston of mammals collected by W. W. Granger Buckhorn, Co., Wyoming ( Long, in 1894, A. Allen included 1965:532) also is M. tJujsanodes. J. (1895a:262) 7Mpus princeps (actually, these were Geomys bursarius lutescens Merriam, Z. Jmdsoniiis) among those mammals 1890.—Long (1965:613) reported a taken in the Black Hills, but not from specimen of the plains pocket gopher the Badlands. Yet, in the species ac- from 0.5 mi N and 1 mi E Beulah, 3550 counts {op. cit. -.266), the only western ft. Crook Co., Wyoming, that allegedly jumping mice obtained by Granger were w as housed in The University of Kansas, listed as from Corral Draw in the Bad- Museum of Natural History. I have ex- lands. I know of no representatives of amined the museum catalogue and spe- this species from western South Dakota cies card files, the field notes and cata- or northeastern Wyoming, where only logues of members of the field party that Z. hudsonius occurs. collected at this site, and all specimens lateralis lateralis of Geomys hursarius from Wyoming in Spermophilus (Say, —The the Museum's collections. Nowhere is 1823). golden-mantled ground does not inhabit the Black Hills there evidence of the indicated specimen squirrel at the time. Coues and Allen and Long's report thus may be consid- present listed a male of this ered as being erroneous. Representatives (1877:818) species that was collected in the "Black Hills" on of this species have been taken on the 21 of 1859 W. S. Wood, who flood plains of the Cheyenne River, on July by the of Lieuten- the southern periphery of the Hills. accompanied expedition ant F. T. the Laramie There are three specimens from 1 mi E Bryan through Mountains, and who, to Edgemont and two specimens from 6 mi my knowledge, did not enter the Black Hills S Hot Springs (UMMZ), Fall River Co., proper. Spermoplidtis lateralis does occur about South Dakota. D. G. Adolphson recently 110 miles to the southeast, in northeast- retrieved several skulls of this species ern from pellets of a barn owl nesting under Albany County, Wyoming (Long, the Hat Creek bridge on Highway 71, 1965:588). 15 mi S Hot Springs, in Fall River County (Martin, 1971a, b). SPECIES OF UNCERTAIN STATUS IN THE BLACK HILLS REGION Phenacomys intermedius intermedius Merriam, 1889.—Cones and Allen (1877: Didelphis marsupialis virginiana Kerr, 216-217) recorded a specimen of Arvic- 1792.—A young opossum that was cap- ola (Pedomys) haydeni (USNM 3056) tured in northern Converse County, Wy- taken from the "Black Hills." They con- oming, by Mr. Hans W. Larsen, subse- sidered the individual as being peculiar quently escaped from a cage in New- and accredited it to being "a young of castle, Weston County, in 1962 (Long, Individuals of this the year." Upon reexamination, J. A. 1965:515). species Allen (1894a:331) assigned the specimen have been observed in recent years in to a new species of Phenacomys (P. fruei, Mellette and Todd counties. South Da- which is in the synonymy of P. interme- kota, and a specimen was captured in dius). In actuality, the indixidual was 1958 at Alliance, Box Butte Co., Ne- collected by W. A. Hammond on 10 Au- braska (Jones, 1964:58). The opossum 150 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY now may be expanding its range into the during the Sangamonian interval and Black Hills from the east by way of the remained in these general areas during deciduous riparian communities of river the Wisconsin age" (Findley, 1955:28). systems. Montane habitat in the Black Hills pre- sumably would be suitable for the dwarf Sorex nanus Merriam, 1895.—The shrew and future application of pitfall dwarf shrew has been reported just east sampling techniques should produce of the Black Hills on the basis of a par- specimens from this region. tially decomposed specimen recovered from a stubblefield 2.5 mi N Fairburn, Myotis grisescens A. H. Howell, 1909. Custer Co., South Dakota. The individ- —On 21 August 1968, Mr. Tim Joseph of ual first was reported as S. vagrans Menden, Missouri, captured a banded a M. (Baker and Findley, 1953:382), but later (B6-02491) male bat (presumably in a small near was assigned to S. nanus (Findley and grisescens) cave Savoy, Lawrence South Baker, 1956:543). The partial skull was Co., Dakota (Gunier, bat had been reexamined recently by Robert S. Hoff- 1971:5). The previously mann and me, and we concur with the captured near Tipton, Moniteau Co., assignment of this specimen to S. nanus. Missouri, in 1967. It was transported to On 3 March 1970, Don Polen, South Higginsville, Lafayette Co., Missouri, Dakota School of Mines and Technol- about 65 miles northwest of the site of to a where it was released ogy, captured a female of this species, capture, place along with two female Sorex cinereus, in a region where there are no known

caves com. . in a grassy field about 10 miles east of ( Gunier, pers. ) the Hills, at a place 3 mi E Boxelder, The intervening distance between Pennington Co., South Dakota (Martin, Higginsville and Savoy is approximately 1971b). External measurements of this 640 linear miles across the semiarid specimen were as follow: total body Northern Great Plains. Presumably, this length, 82; tail length, 35.5; and hind foot environ would be a substantial barrier length, 9. Martin (1971a, b) also re- to a ca\c-inhabiting species such as M. ported a left mandible of S. nanus from grisescens. At no previous time has this a barn owl pellet picked up 15 mi S Hot chiropteran been taken anywhere near Springs, Fall River County. Dwarf South Dakota. The gray bat definitely shrews usually are found in subalpine is not a normal component of the Black and alpine rockslides, at elevations of Hills mammalian fauna and identifica- 8000-10,000 feet; thus, the three speci- tion of bat B6-02491 and the circum- mens from South Dakota are unusual stances of its recapture thus remain sus- both in the low altitude at which they pect. At best, the record represents ab- were taken and in the habitat thev occu- normal wandering of a displaced mam- pied (Hoffmann and Taber, 1960:2.33). mal that was liberated in strange sur- Ecological distribution of the dwarf roundings. shrew seems to be independent of the Euderma maculatum (J. A. Allen, availability of ground water. These ani- 1891).—The rare spotted bat has been mals seldom have been taken in snap- recorded from south-central Montana traps, but recent use of has pitfalls and from Big Horn County in northern proven to be a successful method of ob- Wyoming. This species possibly will be taining specimens (Brown, 1967:622). found along the deep moist canyons that Unfortunately, the of majority trapping are bordered by exposed limestone cliffs in the Hills region has been by use of in the northern portion of the Black and areas have not snap-traps dry rocky Hills. been investigated thoroughly. The ancestor of S. nanus "may have Lepus americanus seclusus Baker and occurred in the Black Hills and isolated Hankins, 1950. —The snowshoe hare evi- mountains of Arizona and New Mexico dentlv does not occur in the Black Hills, TURNER: MAMMALS OF THE BLACK HILLS 151 even though the distribution maps of tenant G. K. Warren, that was collected Burt and Grossenheidcr (1964:212) and by F. V. Hayden in the Black Hills of notations of Seton (1929c:739) indicate Nebraska Territory [now South Dakota] its presence there. Forest Rangers sta- sometime in the period 1857-58. This tioned at Newcastle have "heard of specimen now is housed in the Museum snowshoe rabbits" around Moskee, Crook of Comparative Zoology, Harvard Uni- Co., Wyoming, but I know of no speci- \ersity (MCZ 4817, formerly USNM mens from the Hills. Baker and Hankins 3252/479.5—G. M. Allen, 1931:252), al- (1950:64) stated: "Immediately east- though I ha\'e not examined it. ward of the Bighorns, there are no snow- Ground squirrels of this species pre- shoe rabbits; at least none has been fer sandy soils having sparse or low- reported from any of the higher areas growing vegetation. A single specimen where they might be expected to occur, of S. s. ohsoJetns has been taken along as for example, the Black Hills." Mon- the south fork of White River in south- tane habitats of the Hills presumably western South Dakota (A. H. Howell, would be suitable for this species, how- 1938:131) and 12 others have been re- ever. ported from the three southeasternmost counties in Wyoming, viz. Platte, Lepus californicus melanotis M earns, Goshen, and Laramie (Long, 1965:577). 1890.—Black-tailed jackrabbits rarely Despite extensive collecting, no addi- may occur on the of the Black fringes tional specimens have been obtained in Hills. Game Warden H. J. Brockley and the Black Hills. Several Ranger-Natural- Forest Duke Kocer both have Ranger ists at Wind Cave National Park re- indicated an abundance of blacktails on ported seeing Spermophilus spilosoma the flats to the east and south of the in the upland prairie regions of the Park Hills. I found one killed on a road a in the summer of 1968. Although I sus- few miles southeast of Hot Fall Springs, pect that their identifications were cor- River County, in June 1968. Other speci- rect, I did not observe or take any speci- mens ha\e been reported nearby as fol- mens of this species while conducting low: 1 mi N and 3 mi E Orin, Con\'erse field work within the Park. Co., Wyoming (Long, 1965:552), and 25 mi E and 20 mi S Rapid Citv, Custer Perognathus flavus piperi Goldman, Co., South Dakota (Findley, 1956:2). In 1917.—The silky pocket mouse has been 1956-57, fur dealers from western and taken in northwestern Nebraska, and at southwestern South Dakota reported the type locality 23 mi SW Newcastle, skins of 8500 black-tailed jackrabbits Weston Co., Wyoming (Long, 1965: (South Dakota Conservation Digest, 24: 616). Cones (1875:303) and Cones and 67, June 1957). The majority of these Allen (1877:518) reported a specimen of probably were from Nebraska and Wy- "CricetoclifAis favus' (USNM 3097) oming, but there is evidence of increased from the Black Hills. I have been unable numbers near the Hills. to locate this indi\ idual in order to verify the identity, but know of no other speci- obsoletus Spermophilus spilosoma mens of this species from the region. 1863.—When Kennicott Kennicott, However, the fine sandy loam soils along described (1863:157-158) Spermophilus the South Dakota-Wyoming border pre- he did not a holo- obsoletus, designate sumably would provide a suitable en- but had at hand seven on type, cotypes vironment for P. flavus. which he based the description and name (Lyon and Osgood, 1909:168-169); Reithrodontomys montanus albescens A. H. Howell (1938-130) later selected Gary, 1903. —The plains harvest mouse one of these as the lectotype. Among occurs exclusi\ely in upland habitats, the cotypes is a specimen, received by preferably on sandy soils. The mixed- the Smithsonian Institution from Lieu- grass prairie uplands of Wind Cave 152 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

National Park and some of the drier ern Custer County, South Dakota, and draws of the foothills would seem suit- adjacent Weston County, Wyoming, are able for this species. Although no speci- dominated by sagebrush and mountain mens have been obtained from the Black mahogany that should provide suitable Hills proper, R. montcinus has been taken habitat for Lagunis. Although this mi- from localities near the Hills in Harding, crotine has not been taken there as yet, Meade, Lawrence, and Fall Ri\er coun- the occurrence of preferred habitat in ties. South Dakota, and in Campbell close proximity to the present known and Niobrara counties, Wyoming. For range of the species suggests that the example, on 4 September 1968 I cap- sagebrush vole may inhabit the south- tured a female plains harvest mouse western edge of the Black Hills. the roadside of along grassy Highway 79, Vulpes velox hebes Merriam, 1902. — 1 mi N and 5.5 mi E Hot im- Springs, Of the swift fox, Grinnell (1875:80) mediately adjacent to the Black Hills. wrote that "this pretty little fox is abun- dant everywhere on the plains," and Onychomys leucogaster arcticeps Hoffman (1877:96) indicated that it Hollister, 1914.—A male northern grass- was to be found in the xicinitv of the hopper mouse was obtained at New- Black Hills. Much reduced by hunting, castle, Weston Co., Wyoming on 20 May trapping, and poisoning pressures, this 1894 by Vernon Bailey. Newcastle is fox presumably could still survive in the immediately adjacent to the Black Hills Wind Cave National Park area. The and supports elements of both the Hills species has been reported from Harding fauna and the plains fauna. Additional County in South Dakota (Visher, 1914: specimens have been taken in South Da- 90), southeastern kota on the peripheiy of the Hills at Wyoming (Long, 1965:681), and northwestern Nebraska Buffalo Gap, Custer County; Spring Creek, Pennington County (MHM); 5 (Gary, 1902:67). mi N Hermosa, Pennington County Urocyon cinereoargenteus ocythous on the Pine Indian Res- (SDMT); Ridge Bangs, 1899.—Long (1965:682) noted ervation in nearby Shannon County; and that the westernmost occurrence of this at several localities in Harding County, subspecies is represented b>' a large north of the area. study Specimens of adult female from Owens, Weston Co., from the Moor- Onychomijs leuco^^asier Wyoming (USNM 107892); howexer, I craft Crook area. Co., Wyoming (just have not examined this specimen. Bailey west of the are be- Hills) intergrades (1927:166) indicated that the gray fox tween O. I. and O. I. missouri- arcticeps occurred along the northern border of but are to latter emis, assigned the sub- the Black Hills, and an adult male was species (Long, 1965:638). The northern taken near Deer Ears Butte, Butte Co., mouse occur in the grasshopper may South Dakota, in 1961 (Jones and Hen- of the Red "racetrack" grasslands Valley derson, 1963:283). The "silver gray in the Black Hills. foxes" reported as being "numerous" in the Black Hills area by Dodge (1876: Lagurus curtatus levidensis (Gold- 123) may ha\e belonged to this species. 1941 . — man, ) The sagebrush \'ole occurs west and southwest of the Black Hills Spilogale putorius interrupta (Rafi- in Campbell, Albany, and Laramie coun- nesque, 1820).—Unfortunately, there ties, Wyoming (Long, 1965:660), north- seem to be no specimens of the spotted west of the Hills in Carter and Powder skunk available from the Black Hills River counties, Montana (Hoffmann et region, even though this species un- al, 1969:592), and northeast of the Hills doubtedly occurs there. Merritt Gary in Williams and Morton counties. North related to A. H. Howell (1906:7) that Dakota (Hall and Cockrum, 1953:455). several spotted skunks were killed in the The foothills and canyons of southwest- Black Hills, near Elk Mountain, Custer TURNER: MAMMALS OF THE BLACK HILLS 153

Count)-, in 1902-03. In an unpublislied of North Dakota. It is possible that this manuscript, Gary indicated that this spe- species inhabited the Black Hills prior cies also occurred "near Newcastle" to the advent of European man in the (Long, 1965:702). W. H. Over indi- region. cated in a letter to X. Schantz (1953: Gulo gulo luscus (Linnaeus, 1758).— 125) that "I found a skull of this little The wolverine occurred occasionally in in a cave in the west of mammal slope the Black Hills in the mid-nineteenth the Black Hills." E. R. Hall showed century (Baird, 1858:184; Coues, 1877: of two spotted skunks, photographs 49). The most recently recorded speci- taken at the same in eastern spot Wyo- men for South Dakota is a male killed to R. G. Van Gelder ming, (1959:272); by Mr. Kenneth Long on a sheep ranch one displaxed the pattern of S. p. inter- near (south of) Timber Lake, Dewey and the other resembled S. nipta, p. County, on 10 April 1962 (Lee, 1962:21). these two kinds of <^rcicilis. Thus, spotted Wolverines also have been recorded skunks mav not interbreed also (see from northwestern Wyoming (Long, Mead, It is most desirable 1968:389). 1965:698) and western Nebraska (Jones, tliat be obtained from the specimens 1964:284). Black Hills in the near future in order Lutra canadensis canadensis (Schre- to shed light upon this taxonomic and ber, 1776).—The otter once inhabited at biological problem. Van Gelder (1959: least the river on the the Hills within major systems 251 ) included Black Great Plains, and Grinnell (1875:80) the range of S. p. inferrupta. speculated that it occurred in the Black Martes americana— vulpina (Rafines- Hills. Hoffman (1877:96) encountered que, 1819). Mr. J. Johnson captured occasional skins from west of the Grand a pine marten (USNM 249298) near River Agency. Pringle, Custer Co., South Dakota on 4 Dama dama (Linnaeus, 1758).—The 1930. Whether this specimen January fallow deer is a cervid that has been in- represents an escaped pe*" or was actually troduced from Europe. A herd of fallow indigenous to the Black Hills fauna re- deer range over the Annenburg Ranch mains uncertain. The species may have west of Spearfish, in Crook County, Wy- occurred in the region in the early 1800's; oming. A buck was shot near Hot Hoffman (1877:96) reported seeing sev- Springs, Fall River County, in the au- eral specimens about 8 mi W Grand tumn of 1955 by Hans Palmgreen, and River Indian in Car- Agency, presently another was killed near Piedmont in son South Dakota, and martens County, 1943 (South Dakota Conservation Di- are known from southeastern Wyoming gest, 25:16, July 1956). The irregularity (Long, 1965:690). with which these animals have been Martes pennanti pennanti (Erxleben, taken in the Black Hills suggests that 1777).—The fisher has been taken in the fallow deer has not become firmly northwestern Wyoming (Long, 1965: established in the region, and that these 691), and early trappers reported seeing specimens represent an occasional stray it frequently in the Red River Valley from the Annenburg Ranch.

FACTORS LNFLUENCING DISTRIBUTION AND SPECIATION

MAMMALIAN DISTRIBUTIONAL place the Hills within previously pro- PATTERNS posed biotic distributional areas. Fur- thermore, the diverse topography and Because the Black Hills represent a heterogeneous origins of the mammalian mountainous island habitat surrounded fauna in the Hills (see below) increase the difficulties. the Life- by grass, it is difficult to satisfactorily Nonetheless, 154 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY zone concept of C. H. Merriam (1899) white-tailed jackrabbit, thirteen- lined has some merit, in a general way, when ground squirrel, deer mouse, and mea- applied to the Black Hills. Because life- dow vole) or range freely throughout zones are based on temperature, the both (large and mobile ungulates and zonal boundaries correspond closely with carni\ores, xolant bats, and semiaquatic

has re- furbearers . isotherms. Kendeigh (1954) ) viewed criticisms of this concept. Long (1965:726-729) described sev- Transition Life-zone.—A warmth- eral faunal areas in Wyoming, based on adapted biota occurs on the arid prairie patterns of geographic distribution of uplands and semiarid Red Valley that mammals. In his system, the biota of the surround the Black Hills, in the decidu- Black Hills is associated with the so- ous riparian communities that cross called Great Plains Faunal Area. Its these grasslands, and along the basal divisions of immediate interest here are slopes of the foothills. Steppe-associated briefly discussed below. mammals black-tailed (desert cottontail, The broad arid region that isolates prairie dog, olive-backed pocket mouse, the Black Hills from other forested hispid pocket mouse, Ord's kangaroo rat, mountains in northern and central Wy- prairie \'ole, black-footed ferret, prong- oming comprises the Po\\xler River and and horn, bison) riparian-associated Valley-Bighorn Basin Faunal Di\ision. mammals fox (eastern cottontail, squir- The long-tailed vole and Nuttal's cotton- and white-footed are rel, mouse) fairly tail range westward out of the Hills, limited to the Transition Life-zone in along streamside environs in this region. the Black Hills region. The Cheyenne Plains Faunal Division Canadian Life-zone.—A cold-adapted isolates the Hills from the Laramie biota occupies the forested upper slopes Mountains and other southern mountain of the foothills and the "boreal-cap" ranges in Wyoming. A small tract of (Limestone Plateau, Central Basin, and this unit is in contact ^^'ith the extreme Bear Lodge Mountains) of the Black southwestern corner of the Black Hills, Hills. Se\eral boreal or montane "iso- projecting between the pre\ious and lates" are more or less restricted to this following faunal divisions, and support- life-zone (e.g., red squirrel, northern ing typical plains species. The substrate Hying squirrel, yellow-bellied marmot, of the Sand Hills Faunal Division is red-backed vole, and to an unknown ex- composed of sands or fine sandy loams tent, the ermine). Other species, such as that extend westward from the Sand the least chipmunk, northern pocket Hills of Nebraska, along the South Da- gopher, and long-tailed weasel have en- kota-Wyoming border (Sand Hill Rego- demic in the Canadian Life- subspecies sol Soil Subassociation), and is inhabited zone of the Black whereas the Hills, by various heteromyids and other are- Transition Life-zone is inhabited a by nicolous mammals. different subspecies or represents a The Black Hills Faunal Division is broad zone of intcrgradation between delineated the forested of the races. Some kinds of mammals (the by portion Hills and is inhabited mammals masked shrew, Nuttall's cottontail, by many of boreal or montane affinities. The first bushy-tailed woodrat, long-tailed vole, and meadow jumping mouse) are far three faunal di\isions, and the lower more abundant on the "boreal-cap," than tracts of the fourth, are assignable to the in the Transition Life-zone, where they Transition Life-zone, whereas most of are represented by much lower popula- the fourth faunal division is applicable tion densities. Many species of mammals to the Canadian Life-zone; representa- are not confined to either life-zone, but tive distributions of mammals in these abide in suitable habitats in each (the life-zones were enumerated above. TURNER: MAMMALS OF THE BLACK HILLS 155

ORIGIN OF THE RECENT man, and one (the fox scjuirrel) that MAMMALIAN FAUNA undoubtedly was introduced but prob- OF THE BLACK HILLS ably also reached the Black Hills along natural routes of dispersal. In addition, The present biogeographic analysis 11 species of uncertain status are in- of the Black Hills mammalian fauna is corporated as a component of the Black based on the distributions of 59 indige- Hills fauna; these .species either were re- nous species. These include four species ported from the Hills in early literature, (the wapiti, pronghorn, bison, and or specimens ha\'e been recently ac- mountain sheep) that pre\iousl\- were (juired immediately adjacent to the Hills extirpated and then reintroduced by (Table 29). Three .species (the Norway

Table 29.—Mammals of the Black Hills, listed by faiinal units as discussed in the text. An asterisk denotes species of uncertain status that i^roliably occur (or once did occur) in the Black Hills region.

Widespread Species (26) Sonoran Species (7) Myotis leihii Myotis thysanodes Mijotis hicifugus Sylvilagus auduhonii Lasionyctcris noctivagans "'Lepus californicus Eptcsiciis fiiscus "'Spermophilus spilosoma Lasiiinis cineretis Dipodomys ordii Castor canadensis Reithrodontomys mcgalotis Peromyscus maniculatus "Onychomys leucogaster Ondatra zihethicus

Ercthizon dorsatum Deciduous Forest Species ( 6 ) Canis latrans Myotis keenii Canis lupus Lasiurus borealis Vidpcs vulpes Sylvilagus floridantis Ursus amcricanus Sciurus niger Ursus arctos Peromyscus leucopus " Procyon lotor Urocyon cinereoargenteus Mustcla frcnata Miistela vison Boreomontane Species (10) Taxidca taxtis Sorcx cinereus Mcplntis mej)hitis Tamiasciurus hudsonicus Felis concolor Glaucomys sabrinus Lynx rtiftis Clethrionomys gapperi Cere us canadensis Microtus Odocoileus hemionus pennsylvanicus hudsonius Odocoileus virginianus Zapus "'Martes americana Antilocapra americana Bison bison Mustela erminea "Gulo gulo (10) Steppe Species Lynx canadensis Lepus townsendii tridecemlineatus Spermophilus Cordilleran Species (10) ludovicianus Cynomys * Sorcx nanus Perognath us fasciatus "'Myotis evotis Perognathus hispidus ° volans Reithrodontomys montanus Myotis townsendii Microtus ochrogasier Plecotus ''Vulpes velox Sylvilagus nuttaUii Mustela nigripes Marmota flaviventris "Spilogale putorius Thomomys talpoides (subspecies intcrrupta) Neotoma cincrea Microtus longicaudus Great Basin Species ( 1 ) Eutaniias minimus (karyotype B) Ovis canadensis 156 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY rat, house mouse, and mountain goat) and members of the current plains biota that were introduced from outside of the occupied steppe or savanna conditions study area are not treated in this analy- to the south. Pollen profiles and the pres- sis. ence of relict populations of Microtus 1970 hidovicianus in western Hoffmann and Jones ( ) recently ochrogaster {M. presented a comprehensi\'e biogeo- Louisiana and eastern Texas) and Cyno- graphic analysis of the Recent mam- mys hidovicianus (C. mexicanus in malian fauna of the Northern Great northern Mexico) are evidence of the Plains. Because this thorough study en- southerly displacement of the Great compasses the Black Hills region, much Plains biota to suitable refugia in the of the following account is based on the Full-glacial period (Hoffmann and substance of their report. I also have Jones, 1970:366). Although some re- drawn from my own summary of the stricted grassland may have occurred in vegetation and affinities of the Black the arid rain shadow of the Rocky Moun- flora 26 to 38 and Table tains in it was not Hills ( see pages Late-glacial times, 2). until a shift in atmospheric circulation and climatic e\cnts Steppe Species.—Ten species (14.3%) ensuing occurred, of the Black Hills mammals have evolved in the Pre-Boreal and Boreal periods, that actual of the boreal in response to the semiarid to arid en- replacement vironment of the interior North Ameri- forest by grassland commenced on the Plains. can grasslands and are closely allied Concomitantly, steppe-associated with the Great Plains biota. The distri- mammals dispersed northward, south of the ice and reinhabited the North- butions of Lepiis toicnsendu (listed with front, ern Great Plains and Black Hills both the Great Basin and Campestrian region. faunal units by Armstrong, 1971), As the plains climate became increas- Spermophilus tridecemUneotus, Cijno- ingly warmer and drier, conditions fa- mys hidovicionus, Feroii,naihus fcisciatus, vored eastward extension of the grass- Microtus ocliro^o.ster, Vulpes velox and lands biota. Guilday et al. (1964) re- Mustela nigripes broadly oxerlap the corded two prairie species, Spermophilus Black Hills region, whereas distributions tridecemJineatus and Pedioecetes phasia- of Perognathus hispidus and Reithrodon- neUus (the sharp-tailed grouse), in east- tomys montamis approach their north- ern Pennsylvania in the Late-glacial westernmost limits in the Hills area, and period. Mid-post-glacial time also is the Spilogale putorius (subspecies inier- only period that Spilogale putorius in- reaches its limit. east of the rupta ) westernmost The terrupta ranged Mississippi latter may be distinct from the Great River, into western Illinois (Parmalee Basin race, S. p. gracilis, at the specific and Hoffmeister, 1957). Shifts in the level (Van Gelder, 1959:272; Mead, plains biota presumably occurred many 1968:389). Two additional kinds {An- times and to varying degrees, but most tilocapra americana and Bison bison) recent authors fa\'or the Atlantic period are intimately associated with the prairie as the time of maximal eastward pene- grasslands but range beyond the central tration of the grassland biota. In the steppes and are treated with other eury- Sub-Boreal period, the steppes retreated topic species. Likewise, Leptis californi- to their present limits. and cus, Reithrodontomys megahtis, Differential tolerance to high tempera- Onychomys leucogaster are steppe-asso- tures and accompanying heat stress may ciated mammals but have a much more have dictated the post-glacial moxements extensive distribution in the Southwest of some members of the plains biota. and Mexico. adjacent For example, Microtus ochrogaster is During the maximal Wisconsin gla- much more tolerant of xeric conditions ciation, much of the Northern Great than is its boreomontane congener, xA/. Plains was covered by a boreal forest pennsylvanicus; yet, populations of M. TURNER: MAMMALS OF THE BLACK HILLS 157

ha\c decreased ochrogaster drastically are associated with the interior grass- in recent droughts ( Wooster, 1935, 1939; lands but have extensive distributions Gier, 1967). The easternmost margins beyond the central steppes. Lepu.s cali- of the plains-grasslands presumabK' foniicu.s, Di])odomys ordii, Reithrodonto- would be somewhat more mesie, and the mys me'^alotis and Onychomys leuco- eastward shift in A/. ocJiwgaster dining gcister also range into the Great Basin, as drier post-glacial times may haxe been does Myotic thysanodes. The ranges of in response to decreased evaporative Syhilagus- auduhonii and Spennophihis stress in such areas. Con\'ersely, the oc- spdosovui are more widespread west of currence of Pero!j.natlius Jiispidus in east- the Black Hills, but rarely exceed 40° ern Missouri in Late-glacial or early latitude in the far west. Leptis ccdiforni- Holocene time (Oesch, 1967) may be cus, R. megalotk, and O. leticogasier credited to the ability of this heteromyid also are apportioned far east of the Hills, to tolerate extremely cold temperatures as was S. auduhonii during chronic (Jones, 1964:173). As a final example droughts of a few decades ago (Hoff^- of temperature-influenced shifts in dis- mann and Jones, 1970:382). tribution, the present northward retreat Both L. californicus and S. spilosoma of Lepiis tounsendii from the southern approximate their northernmost limits part of its range and its current expan- on the Great Plains along the south- sion to the north and northeast may be eastern margin of the Black Hills. The correlated with the gradual warming former species has extended northward trend now in progress on the North into South Dakota within historic time, American continent (op. cif.: 113). whereas the latter has not been taken in the Hills since the of 1857-58. Other factors influencing past distri- period butions of steppe mammals include the Myotis thysanodes is represented by an endemic in the Black presence of efl^ecti\'e barriers to dispersal population Hills and the availability of food sources. The region (subspecies polia.sapen.sis). Missouri River limited the eastward ex- Either a few individuals managed to pansion of P. hisvidus and Reithrodonto- cross intervening barriers to initiate a the )nys montanus. Past and present ranges disjunct colony, or population in the Hills is a relict that resulted from the of Musiela nifiripes correspond closely to those of Cynomys ludovicianns. There fragmentation of a formerly more wide- are some exceptions to the last men- spread distribution. tioned example. For instance, M. ni- Great Rasin Species.—Although many iiripes is represented in late Pleistocene members of other faunal units range deposits in Jaguar Cave in Idaho, and in into the Great Basin (Armstrong, 1971, Orr Cave in Montana, but remains of listed Myotis volan.s, Lepus townsendii, Cynomys apparently were not present in TJiomoinys talpoides and Antdocapra either deposit (Guilday and Adam, americana as components of this faunal 1967:30). unit), only one species (1.4%) of the Black Hills Sonoran Species. —Seven kinds of mammalian fauna presum- mammals (10.0%) present in the Black ably originated there. Hoffmann and a Great Hills are typical of the Chihuahuan- Jones (1970:383-385) proposed Sonoran Region of southwestern United Basin-Great Plains pattern of differen- tiation for Eutamias States and adjacent Mexican Plateau. niinimus, and indi- cated that of Like the steppe species, these mammals subspeciation Thomomys and evidently originated under the dictate of talpoides Spdogale putorius also an arid climatic regime, and subse- may fit this general pattern. Definitive evidence quently invaded the Hills region in times concerning differentiation of the latter two is as of post-glacial warmth and dryness, species lacking yet. prol^ably in the late Boreal and early Recent analysis of chromosomal Atlantic periods. Most of these species structure of Eutamias minimus (Sutton 158 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY and Nadler, 1969:526) indicates diat of the early Holocene, or possibly of boreal forest races horeaJis and ( neglec- prior interglacial periods, permitted tus), Rocky Mountain subspecies (ari- movement of Eutamias minimus across zonemis and consohrinus), the Laramie the W\'oming Basin, onto the Northern Mountains race (operarius), and two Great Plains. A shift of the upper-air secondarily steppe-adapted subspecies antic>X'lonic eddy northeastward from (niininuis of the Wyoming Basin and the Great Basin in the Sub-Atlantic period of the San Luis and 1967 have canji Valley) possess ( Bryson Wendland, ) may karyotype "A." The Great Basin races influenced dispersal of the biota at a {scrutator and pictus). Northern Great later time. The boreal forest chipmunks Plains subspecies (palliclus and cacode- pre\iously in the plains region, theoreti- ?nus), and two montane populations cally retreated upslope into montane en- (confinis in the Bighorn Mountains and vironments, coincident with retreat of silvatictis in the Black Hills) possess boreal elements from the prairie. Ap- karyotype "B." The absence of hetero- parently freed from intense competition, morphic chromosomes is evidence that the invaders from the Great Basin then subspecies with these two different kary- adapted to the central steppes environ- otypes may in fact represent sibling ment. species (Sutton and Nadler, 1969). That Eutamias minimus confinis and E. m. silvaticus to have been During maximal glaciation, the an- appear cestral stock of the Great Plains races derived from adjacent plains popula- tions rather than from boreal forest or of Eutamias minimus evidently became Mountain ancestors is indicati\'e disjunct from topical boreal representa- Rocky tives. Isolated by a barrier of coniferous of several periods of isolation as de- forest that intervened between the Great scribed above. Otherwise, the montane Basin and Wyoming Basin, the ancestral derivatives of the plains populations would ha\e met intense com- stock e\'ol\'ed in response to arid steppe extremely and semidesert enxirons that were asso- petition from residual boreal chipmunks, the ciated with sagebrush-grass communi- already occupying Bighorn Moun- tains Hills. ties, as well as coniferous ^^'oodlands. The and Black Probably, boreal- races would have a interposing coniferous forest barrier pre- adapted competitive sumably was intermittent, and formed advantage in a montane environment in response to climatic conditions that over those that were steppe-adapted. fa\'ored each continental and Cordilleran Under what conditions steppe-adapted races would find these montane areas glaciation; thus, it was fashioned by free of either southerly displacement of boreal boreal-adapted chipmunks, or forests or by descent of montane forest. would become competitively superior to the remains For example, regional montane biotic latter, problematical.— zones throughout the cordillera were de- Deciduous Forest Species. Six spe- cies of the Black Hills pressed vertically 4000-4500 feet, con- (8.6%) mammalian current with a decrease of 16-17 F in fauna have affinities with eastern hard- wood mesic or summer temperature in the Full-glacial forests, grasslands, prairie period (Richmond, 1965:228, and oth- riparian communities. Most of these ers). A similar descent of the lower tree- mammals reach the westernmost limits line evidently occurred in the Sub-Boreal of their continuous range in the xicinity the Hills. and possibly other periods. of As the boreal forest retreated north- Mijotis keenii (subspecies septen- ward and the montane woodland as- trionalis) and Sciurus niger are restricted cended upslope, a relatively low non- to temperate eastern North America, forested pass opened across the Conti- whereas Lasiurus horcalis (a seasonal nental Dixide between the Great Basin migrant in the Hills region), Sylvilagus and Wyoming Basin. Thus the climate jioridanus, Feromijscus leucopus and TURNER: MAMMALS OF THE BEACK IIILES 159

Urocijon cinereoargenteiis aic> widely the Black Hills area within historic time. distributed in the temperate eastern and WUles-pread Species.—Twenty-six subtropical southern portions of the con- species (37.1%) of the Black Hills mam- tinent. The last nientioncxl sp(>ci(\s orig- mals are eurytopic, with no apparent inated in the Neotropical Region, but faunal affinities in relation to the Hills invaded the Northern Great Plains from region. These kinds are present in the the east (Hoffmann and Jones, 1970: study area because they typically are

377 . Whether keenii is mobile. ) Mijotis an actual extremely They are euryecious "relict" or a seasonal migrant to the with a wide range of tolerance for vari- Black Hills is still in question. Peroniys- ous environmental factors, or are sten- cus leucopus presently occurs on the oecious but encounter specific environ- Northern Great Plains in scattered lo- mental requisites in several segregated calities that are relatively isolated from habitats throughout the region. their respective contiguous ranges (see Volant bats, and large ungulates and Figs. 15 and 16, Hoffmann and Jones, carnivores, are exceedingly mobile, rang- 1970:38(8). The currently disjunct popu- ing over vast areas that may be com- lations of P. leucopus may represent iso- posed of several different environs. Of lated segments of a formerly more or less the eurytopic chiropteran fauna, Myotis continuous and interbreeding population leibii is restricted to temperate North that dispersed northwestward behind America, whereas M. lucifugiis and the retreating ice sheet and was subse- Lasionycteris noctivagans also range (juently extirpated in the more xeric cli- over boreal portions of the continent, matic regimes. Alternatively, they may and Eptesicus fiisctis and Lariurus cine- typify the disruption of a continuous reiis additionally are distributed to the dendritic distribution that paralleled Neotropical Region. Two of these spe- mesic drainage systems westward across cies (L. noctivagans and L. cinereus) the prairies. The latter explanation is are seasonal migrants in the Black Hills. favored most recent authors. Cur- by Of the large game mammals and fur- between these limited rently, gene-flow bearers whose distributions overlap the is reduced or and populations absent, Black Hills, five are primarily temperate each presumably is adapting independ- kinds Taxidea ( taxiis, Lynx rufus, Odo- to its immediate en\'ironmental ently coileus hemionus, Antilocapra americana as evidenced the rela- conditions, by and Bison bison), three are mainly tem- small and dark mice on the Black tively perate-boreal taxa (Vidpes vulpes, Me- Hills. phitis mephitis and Cervus canadensis), Elements of the eastern deciduous three are temperate-boreal-subtropical forest replaced boreal forest species types (Canis hipus, Ursus americanus, along the eastern periphery of the North- and U. arctos) and five species (Canis ern Great Plains in Pre-Boreal times. latrons, Procyon lotor, Miistela frenata, Members of this faunal unit presumably Felis concohr, and Odocoileus virgini- then dispersed westward along wooded ana) range from temperate regions into tributaries of the Missouri River system, both boreal and Neotropical areas. Three to be excluded from the of these U. P. only plains (or mammals ( americanus, isolated in suitable refugia) in the sub- lotor, and O. virginiana) are primarily sequent arid Atlantic time. With ameli- associated with woodlands throughout oration of climatic conditions in the their ranges, whereas four others (T. more mesic Sub-Atlantic and Neo-At- taxiis, O. hemionus, A. americana, and lantic periods, eastern species could dis- B. bison) are associated for the most perse westward along gallery forests part with nonforested environs. The re- once more. Several species (Sciurus maining nine species occur in both major niger and Urocyon cinereoargenteus) habitats. have extended their ranges westward to Three stenoecious aquatic or semi- 160 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

are aquatic mammals included in the northward (Hoffmann and Jones, 1970: eurytopic faunal unit, Castor canadensis. 386 and Fig. 1). Ondatra zihethicus, and Mustela vison. Relict populations of northern mam- These are temperate-boreal species with mals that now are disjunct from other specialized niche requirements that are such populations, and from the main fulfilled by a variety of aquatic habitats range of the species, are an indication throughout their range. of the widespread extirpation of boreal Among widespread terrestrial ro- elements during the Atlantic period and other dents, Erethizon dorsaitim is a temper- warm, dry climatic episodes. Only the hardiest ate-boreal species that usually is asso- colonies survived the xeric ciated with coniferous trees, but has an conditions, and these abided in the fa- extensive distribution beyond such for- vorable or tolerable cool and mesic pock- ests. Peromyscus manlculatus best ex- ets on the plains, or in montane regions. a Three boreomontane mammals Sorex emplifies euryecious taxon. This species ( Microtus consists of a long series of interbreeding, cinereus, pennsylvanicus and or potentially interbreeding, populations Zapiis hudsonius) are widely distrib- distributed from northern boreal North uted across the Northern Great Plains in America southward to Oaxaca, Mexico. suitable riparian communities. The lat- It is ubiquitous, occupying a remarkable ter species lacks a western montane distribution variety of habitats and, being quite re- typical of other members of sponsive to local environmental selective this faunal unit, and often occurs in pressures, has formed numerous eco- populations that are isolated from oth- types throughout its range. ers by broad expanses of inhospitable terrain. Boreomontane Species.—Ten species few (14.3%) of the mammals inhabiting the A species (Martes americana, Black Hills are distributed both in the Gulo gulo and Lynx canadensis) repre- northern boreal forest and in the conif- sent northern kinds that presumably in- erous forest of western montane areas. habited the Black Hills in the recent past, Mammals of the boreomontane faunal but ha\'e since become extinct there. unit for the most part range much far- Occasionally, some representatives of ther north than do members of the cor- these species may still wander into the dilleran faunal unit; southern limits of Hills region. distribution also farther north average The remaining four species (Tamia- for the former than for the latter. sciurtis hudsonicus, Glmicomys sahrinns, Boreal forest, displaced southward Clethrionomys ii^apperi, and Mustela er- by continental glaciation, and montane minea) represent "glacial relicts" that forest, displaced downslope, eastward probably were isolated subsequent to and the southward by Cordilleran glacia- northerly retreat of the ice sheet. tion, probably intermixed along a broad Mustela erminea is disjunct in the Black front, maintaining a continuous conifer- Hills to an unknown extent; it is present ous woodland throughout the Powder in the Laramie Mountains, but appar- River Valley and Cheyenne Rixer plains. ently absent in the Bighorn Mountains. The boreomontane biota in the Black The level of divergence of Clethriono- Hills was thereby connected with that mys gapperi ])revicaudus (previously as a of the Bighorn and Laramie mountains regarded distinct species) suggests that this until at \'o]e may ha\'e become disjunct least the Pre-Boreal period. Sev- earlier than did other "isolates," or may eral forested corridors that provided have diverged at a faster rate; the op- avenues for dispersal of boreomontane posite supposition may apply to Glau- mammals remained on escarpments and comys sabrinus and M. erminea (see the other topographic breaks well after the following section on Speciation). Cock- coniferous forest retreated upslope and rum and Fitch (1952) indicated that TURNER: MAMMALS OK THE BLACK HILLS 161

C. l)revicau(lus is related to slides in affinities. However, until loniiicaiidus hav(> relatively broad niches, more evidence is a\ ailable, my comments dispersing westward across the Powder here are based on the more northern Ri\'er Valley plains in association with distribution of the species as a unit. riparian communities. Of the cordilleran The subspecies of T. hudsonicus in faunal unit, only Marniota flaviventris the Black Hills (cJakotensis) appears to represents a distinctly disjunct popula- be more closely related to the reddish- tion. Isolation of this marmot in the colored boreal forest populations than Black Hills presumably occurred in the to the darker-colored Rocky Mountain manner assumed for other "glacial re- races (Hoffmann and Jones, 1970:372). licts." Reddish-colored squirrels in the north- Several taxa present in the Bighorn ern Laramie Mountains, southeastern and Laramie mountains either failed to Montana, northwestern South Dakota, disperse to the Black Hills, or were extir- and other outlying areas probably are pated there during xeric climatic regimes remnants of the former population that (Hoffmann and Jones, 1970:376 and remain distributed along past corridors Table 2). This fact, in conjunction with and dispersal routes. Assuming that da- the presence of other taxa in the Big- kotensis originated in response to selec- horn Mountains that are absent in the ti\e pressures of the Black Hills environ- Laramie Mountains, and vice versa, sug- ment, the race either enjoyed a broader gests a filtering effect of past barriers, or distribution in early Holocene time, or differential survival of species reaching has since dispersed outward from the these montane environments, or both. Hills via pine-filled canyon systems and Using the distribution maps of Long conifer-dominated escarpments. (1965), the number of subspecies of Cordilleran Species.—Ten species mammals in each of the three indicated (14.3%) of the Black Hills mammalian montane environs was ascertained. fauna represent elements that were dis- Twenty-five taxa occur in all three placed into the region by Cordilleran ranges, and 19 kinds are common to the ice sheets. These mammals evidently Black Hills and Laramie Mountains, but became stranded after retreating upslope do not abide in the Bighorns. Four sub- in response to increasing aridity in post- species are shared by the Black Hills and glacial time. All but three species attain Bighorn Mountains, but not by the Lara- their easternmost limits on the Northern mie Mountains. Eleven races are com- Great Plains in the Black Hills—the mon to the Bighorn and Laramie moun- range of Neotoma cinerea and Ov)is cana- tains, but do not occur in the Black Hills. densis extend slightly beyond, terminat- Sixteen kinds inhabit the Black Hills, ing in western South Dakota, and Thom- but are absent from the other two ranges. omijs tolpoides is broadly distributed Eleven subspecies occur in the Bighorn eastward to approximately 98 longi- Mountains, but not in the Black Hills or tude. The plains and montane races of Laramie Mountains, and 13 subspecies the latter species may have a pattern of live in the Laramie Mountains, but not differentiation similar to that of Eutani- in the Black Hills or Bighorns. Three ias minimus. species (Etitamias minimus, Marmota and Myotis evotis, M. vohns, Plecotus fiaviventris, Thomomijs talpoides) have diversified into toicnsendii, 'Neotoma cinerea and Ovis separate subspecies in each of the aforementioned mountain- canadensis generally are associated with ous montane forests, rocky outcrops, or bad- regions. lands topography in western North Employing these data, a percentage America, whereas Sorex nanus usually is index of faunal resemblance (Simpson, associated with subalpine or alpine rock 1943) can be calculated for each pair 162 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY of montane areas. Such indices are not dilleran kinds, the Laramie Mountains the Black Hills absolute, but are meaningful only when fauna resembles that of comparing neighboring faunas, thereby due to a greater presence of steppe, So- and eastern The Black yielding a relati\e measure of similarity noran, species. Hills and on the (Udvardy, 1969:273). The larger the Bighorn Mountains, both exceed the Laramie percentage index, the more closely the other hand, Mountains in relative of faunal pairs resemble each other. Com- proportions and mam- puted indices between the various pairs boreomontane widespread are as follow: Black Hills and Bighorn mals. The latter mountainous region is in ele\ation and Mountains, 56.9; Black Hills and Lara- intermediate ( perhaps as com- mie Mountains, 68.7; Bighorn and Lara- other environmental factors) mie mountains, 70.6. pared to the former two montane areas. Another avenue of exchange Biogeographic analysis of affinities possible between the Black Hills and of the mammals in the Bighorn and Lara- Bighorn ha\'e occurred to the mie mountains were undertaken in a Mountains may via the wooded manner similar to that used in the analy- north, Rosebud-Tongue river breaks and conifer-clad Pine sis of the Black Hills biota. Percentages Long Hills of southeastern Montana and of the mammalian fauna attributable to northwestern South Dakota. the respective faunal units are given for the Black Hills, followed sequentially by that of the Bighorn Mountains and Lara- SPECIATION AND GEOGRAPHIC mie Mountains as follow: widespread VARIATION species, 37.1, 41.2, 36.8; steppe species, 14.3, 9.8, 11.8; Great Basin species, 1.4, BioticaUy-induced Selection Pres- 2.0, 2.9; Sonoran species, 10.0, 7.8, 8.8; sures.—Two pairs of related species deciduous forest species, 8.6, 0.0, 4.4; {Mijotis evotis and M. thysanodes; Syl- boreomontane species, 14.3, 13.7, 11.8; vihgiis floridanus and S. nuttallii) are cordilleran species, 14.3, 25.5, 23.5. parapatric in the Black Hills region. Indices of faunal resemblance and Several other pairs of species {Sorex biogeographic analyses both suggest that cinereiis and S. nanus; Sylvilagus audti- coniferous forest corridors between the honii and S. floridanus; Reithrodonio- Bighorn Mountains and Black Hills were mys megalotis and R. montanus; Pero- indirect—\'ia the northern Laramie myscus leucopus and P. maniculatus; ranges (Casper and Haystacks), the Microtus longicaudiis and M. pennsyl- Hartville Uplift, and the Pine Ridge vanicus; M. ochrogaster and M. penn- escarpment (Hoffmann and Jones, 1970: sylvanicus; Mustela erminea and M.

386 . This would account for the hemionus and O. ) greater frenata; Odocodeus faunal similarities between the Bighorn virginiana) are known to be sympatric, and Laramie mountains, and between but ecologically distinct to varying or the Laramie Mountains and Black Hills. unkno\\'n degrees. Specific distinctness The relatively high percentage of cor- of such species pairs strongly suggests dilleran mammals in the two former extrinsic isolation in the past and subse- montane environs, as compared to the quent development of intrinsic isolating Hills, further implies that the indirect mechanisms in response to differential connection was of greater duration, or selection pressures. Indeed, with the was a much better dispersal route, or exception of the eurytopic deer, each both, than was a direct extension of member of the aforementioned species coniferous forest eastward from the Big- pairs belongs to a separate faunal unit horn Mountains to the Hills. of different biogeographic affinity (see Whereas the mammalian faunas of Table 29). Interaction among closely the Bighorn and Laramie mountains are related species favors differential ex- similar due to a predominance of cor- ploitation of similar niches, thereby de- TURNER: MAMMALS OF THE BLACK HILLS 163

the of in creasing intensity interspecific suggests situ divergence of clinally competition. connected ecotypes in the Black Hills Seemingly, selection pressures would region. be less stringent on those species that do For the most part, the Black Hills not compete with congeners in the study are inhabited by races of mammals that also inhabit area. However, interspecific and ecologi- the encompassing plains. cal interactions between various sym- Continual exchange of hereditary ma- terials between the Hills and patric genera ( Lepus and Sylvilagus; plains Eufamias and SpenuopJiihis; GJaucomys, populations seems probable. Dichroma- Sciiinis and Tainiasciunis; Cletliriono- tism (dark and pale phenotypes) and mys and Micwtiis), among various chi- extreme individual variation in the Black ropterans, and among several heteromy- Hills populations of Peromyscus manicu- ids may favor partitioning of similar latus- (Osgood, 1909:78; Dice, 1942:1- niches in the Hills region due to these 10), and of Spernwphilus tridecemline- atus and biotically-induced selection pressures. Microtus longicaudus (Long, are due to dis- EnvironmenfaUy-indiiced Selection 1965:738) presumably of the Pressures.—Many kinds of mammals ruption strong selection for darker coloration in the Hills influx have differentiated into more or less dis- humid by of alleles from forms on the arid tinct subspecies as the result of selection paler pressures engendered by the Black Hills plains. environment. Four of these subspecies An additional source of variability in mammals within the Hills is due to the ( Myotis thysanodes pahasapensis, Ta- miasciunis hudsonicus dacotetisis, Mar- great diversity in physiognomy, exposure of nwfa flaviventris dacoto, and Clethriono- soils, and other general edaphic char- acteristics. small mys gapperi brevicaudus) currently are Because mammals are disjunct and continue to adapt inde- intimately associated with their habitats, pendently. Three others (Eutomias selection pressures in one area may be mininuis silvaticiis, Thomomys talpoides quite different from those in another. nelndosns and Miistelo frenata olleni) For example, the relatively dry, hard- are represented on the surrounding wood-dominated drainage system of Big plains by different subspecies (E. m. Spearfish Canyon and similar streambeds pallidus, T. t. hidhtus and T. t. pierreic- in the Bear Lodge Mountains contrast neva- olus, M. f. longicouda and M. f. sharply with the cool, mesic Little Spear- derisis), and are presumed to be in ge- fish Canyon that is characterized by lux- netic contact with these surrounding uriant moist meadows and spruce-bor- populations. The presence of parapatric dered streams. As indicated in Figure 12, subspecies implies either that the two the general climatic regime of the north- races formerly were isolated and di- ern Hills differs from that of the southern verged (but not to the extent of devel- Hills. Additionally, the climatic and en- oping intrinsic isolating mechanisms) in vironmental gradations from relatively response to different environments be- mesic higher elevations, to semiarid fore subsequent contact with the step foothills, to arid plains is much more cline, or that subspecific divergence oc- gradual in the southern Hills as com- curred in situ due to strong selective pared to the abrupt changes typical of pressures that formed separate clinally the northern Hills (see page 112). The connected ecotypes. The extreme con- occurrence of Eutamias minimus polli- trast between the Black Hills and Great dus, rather than E. m. silvaticus in the Plains in climate, topography, and bi- extreme southern and southeastern Black otic communities enhances the feasibility Hills probably is a reflection of the grad- of the latter mechanism. The general ual gradation of environmental factors, pattern of \'ariation in other mammalian hence, of environmentally-induced selec- species (see following discussion) also tion pressures. Other small mammals in 164 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY the southern portion of the Black Hills netic drift may also play a role in in- generally tend to be slightly paler than creasing recessive homozygosity, thereby those of the northern area. In addition, enhancing differentiation of these dis- the dorsal pelage of several mammals in junct populations. reflect red the northwestern sections The broad arid regions (Belle hues more intensely than those from Fourche, Cheyenne, and Powder river that other areas. Species \'ary noticeably plains) that isolate the Hills from other within the Black Hills include Eutamias montane areas are inadequate barriers Tamiasciurus Tho- minimus, hudsonicus, to some species. These rivers provide manicu- momys talpoides, Peromyscus routes of immigration and emigration Microtus latus, Neotoma cinerea and for Sorex cinereus, Sylvilagus floridanus pennsylvaniciis. (to the north, south and east), Sylvila- The major effect of selection in the gus nuttoUii (to the west), Sciurus niger, Black Hills as a unit seems to be in Thomomys talpoides, Peromyscus leuco- accord with Gloger's Rule, producing pus, Microtus longicaudus, Microtus relatively dark and reddish-colored pop- pennsylvanicus, Zapus liudsonius, and ulations. The positive correlation in- various semiaquatic mammals. Thus, dicated by Stebler (1939) and Dice genetic flow is probable along streams in (1942) between the color of dorsal these species, between riparian-resti'icted pelage of mammals and the color of plains populations and those on the exposed soils in the Black Hills and Black Hills. Long (1965:733) noted that on the Great Plains already has been dis- among lagomorphs, the species most de- cussed (see page 99). Dice (1947:19), pendent on riparian habitats is darkest after undertaking several experiments in color—Sylvilagus floridanus (dark), that established significant selection by S. tiuttallii (intermediate), and S. audu- barn owls for deer mice contrasting in bonii (pale). coloration with their background, con- The degree of differentiation of dis- cluded that ". . . natural selection can junct species in the Black Hills may be theoretically produce rapid evolution used as an indirect measure of the dura- whenever a genetically variable popula- tion of such isolation. Such indications tion is to its action." exposed Popula- are relative and may be modified greatly tions presumably respond to diff^erent by unknown differential rates of evolu- environments variation in through gene tion of the species in question. For ex- frequencies, but natural selection tends ample, Clethrionomys gapperi hrevi- to minimize the variability, increasing caudus differs from C. g. galei of the frequency of adaptive phenotypes and Rocky Mountains in 11 mensural param- thus local creating ecotypes. eters ( see also Cockrum and Fitch, 1952 ) Clark (1938) indicated that the pale and previously was treated as a distinct ochraceous-buff color of deer mice on species (Bailey, 1897:129). The remain- the plains is a character dominant over ing subspccific isolates (Myotis thysa- the recessive darker coloration that fre- nodes, Tamiasciurus hudsonicus, and quently occurs in deer mice inhabiting Marmota flaviventris) differ from the montane areas. Seemingly, the environ- nearest conspecific populations mainly in ment in the Black Hills favors a higher color, or differ in mensural characters to frequency of recessive dark alleles, but a comparatively lesser extent than en- the gene pool is disrupted by the influx countered between the two subspecies of dominant pale alleles from the sur- of Clethrionomys gapperi. Myotis keenii, rounding plains populations, resulting Glaucomys sahrinus, and Peromyscus in dichromatism within Hills popula- leucopus also are isolated to differing tions. If a similar genetic scheme may degrees and currently appear to be di- be assumed for the several isolated mam- verging under thc^ dictate of natural se- mals in the Black Hills region, then ge- lection in the Black Hills. The Hills TURNER: MAMMALS OF TJIE BLACK HILLS 165 population of Mustela erminea appears pliilus tridecemlineatus, Neotoma cin- disjunct, but cnidcntly does not differ erea, Peromyscus maniculatus, Microtus notabh' from members of this species in longicaudtis, Microtus penn.sylvanicus the Laramie Mountains. Hence, C. gap- and Microtus ochrogaster all seem to be jicri either became isolated in the Black evolving darker color in the study area. Hills at an earlier time than did the On the other hand, representatives of other "glacial relicts," or has diverged Reitlirodontomys megalotis that occur at a faster rate due to a shorter genera- in the Hills show no appreciable change tion time. Conversely, those species that in coloration when compared to those ha\e differentiated in the Hills region of the surrounding plains. Finally, tliree to a lesser extent either were isolated in species (Sorex cinereus, Peromyscus a later time period or ha\e responded moniculotus, and Zapus hudsonicus) more slowly to selection pressure in- vary clinally in coloration, and in exter- curred by the Black Hills environment. nal and cranial dimensions northwest- kinds of with across the Plains Manv mammals broad wardly Great ( Dice, distributions that oxerlap the Black Hills 1941:16; Robertson, 1971). With excep- have not diverged subspecifically, but tion of somewhat darker color, the Black display some modification in coloration Hills apparently do not interrupt the within the Hills. Populations of Spermo- clinal nature of variation in these species.

SUMMARY

The Black Hills of western South miles to the southwest and west, respec- Dakota and northeastern Wyoming con- tively, in Wyoming. stitute an elliptically-shaped, isolated This report delimits and describes mountainous region of approximately the Black Hills as a natural zoogeo- miles that resulted from 4000 square graphic unit, considers the autecology intermittent domal uplifts in Cretaceous, and general distribution of each mam- times. En- Miocene, and Pleistocene malian species in the Hills, discusses the tirely surrounded by the non-glaciated geographic variation and inferred specia- Missouri Plateau section of the Northern tion of these mammals, and analyzes the Great Plains physiographic province, and interprets the probable biogeo- 4000 Hills rise above the plains about graphic affinities of various species in 3000 feet on feet on the east and some light of conjectured changes in late the west; the highest point is 7242 feet Pleistocene and Holocene environments. above sea level. Most of the area lies Six orders and 19 families, including 44 of the within the drainages Cheyenne genera and 62 species, comprise the and Belle Fourche rivers. As here con- Black Hills mammalian fauna. the Black Hills are delimited sidered, Biogeographical analysis of the Recent the distribution of shale and by Jurassic mammalian species of the Black Hills sandstone of the Sundance Formation. suggests the following faunal affinities: The thus circumscribed includes region 37.1 percent mdespread, 14.3 percent the Black Hills as well as the proper, steppe, 14.3 percent boreomontane, 14.3 Bear Mountains and Devils Tower Lodge percent cordilleran, 10.0 percent So- of Wyoming; these tracts are closely noran, 8.6 percent deciduous forest, and allied geographically, geologically, cli- 1.4 percent Great Basin. Indices of and biologi- matically, physiognomically, and nearest to faunal resemblance biogeographic cally. Ranges of mountains that coniferous forest the Black Hills are the Laramie Moun- analyses suggest corridors between the Moun- tains (to 10,272 ft) and Big Horn Moun- Bighorn tains and Black Hills in the Pleistocene tains (to 13,165 ft), approximately 150 166 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

time were indirect; this accounts for the niches due to these biotically-induced faunal similarities between the Bighorn selection pressures. and Laramie mountains, and between Four endemic subspecies currently the Laramie Mountains and Black Hills. are disjunct from contiguous popula- The relatively high percentage of cor- tions, whereas three others are in genetic dilleran mammals in the two montane contact with different subspecies on the i environs, as compared to the Hills, fur- surrounding plains. Such parapatric sub- 1 ther implies that the indirect connection species probably di\'erged in situ due to was of greater duration, or was a much local selection pressures that formed better route or both than was dispersal ( ) separate, clinally-connected ecotypes. a direct extension of coniferous forest Most races in the Hills also inhabit eastward from the Bighorn Mountains the plains. Dichromatism in several spe- to the Hills. cies is due to disruption of selection for darker coloration in the Interaction among closely related spe- humid Hills cies favors differential exploitation of by influx of pale alleles from arid plains similar niches, thereby decreasing the populations. Small mammals in the southern intensity of interspecific competition. Hills tend to be slightly paler than in Ten pairs of related species in the Hills those the northwestern Hills, which region either are parapatric or sym- reflect red hues more intensely than patric but ecologically distinct to varying individuals from other areas. degrees. Specific distinctness of such Many eurytopic mammals have not species pairs strongly suggests extrinsic diverged subspecifically in the Hills, but isolation in the past and subsequent display modification in coloration; one development of intrinsic isolated mech- species shows no apparent change. Three anisms in response to differential selec- species vary clinally northwestwardly tion pressures. Intergeneric competition across the plains; the Black Hills do not also may favor partitioning of similar interrupt these clines.

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White, E. M. 1939. The effects of drouth on rodent popu- 1961. Drainage alignment of western South lations. Turtox News, 17:26-27. Dakota. Amer. Jour. Sci., 259:207- Wright, H.E.,jR. 210. 1968. History of the Prairie Penninsula. Pp. White, J. A. 78-88, in The Quaternary of Illinois

1952. A new cliipmunk (genus Eutatuias) (R. E. Bergstrom, ed. ), Spec. Publ. from the Black Hills. Univ. Kansas Univ. Illinois Coll. Agric, 14:1-179. Publ., Mus. Nat. Hist., 5:259-262. 1970. Vegetational liistory of the Central 1953a. Genera and subgenera of chipmunks. Plains. Pp. 157-172, in Pleistocene Univ. Kansas Publ., Mus. Nat. Hist., and Recent environments of the Cen-

5:543-561. tral Great Plains (W. Dort, Jr., and

1953b. Geographic distribution and taxonomy J. K. Jones, Jr., eds. ), Univ. Press of the chipmunks of Wyoming. Univ. Kansas, Lawrence, 433 pp. Kansas Mus. Nat. 5:583- Publ., Hist., Young, S .P. 610. 1944 Their history, life habits, economic R. L. Wilson, status, and control. Pp. 1-386, in The 1967. The Pleistocene vertebrates of Michi- Wolves of North America (S. P. Acad. gan. Papers Michigan Sci., Arts, Young and E. A. Goldman), Amer. and Letters, 52:197-234. Wildlife Inst., Washington, D. C, WiNCHELL, N. H. XX-}- 636 pp. 1875. Geological report. Pp. 21-66, in Re- 1946a. The wolf in North American history. port of a reconnaissance of the Black The Caxton Printers, Ltd., Caldwell, Hills of Dakota made in tlie summer Idaho, 149 pp. of 1874 (W. Ludlow), Dept. Engi- 1946b, life economic neers, U. S. Army, 121 pp. History, habits, status, Woodward, H. R. and control. Pp. 1-173, in The puma, 1930. Abnormal growth of incisors in the mysterious American cat (S. P. Young Fiber zihethicus. Proc. South Dakota and E. A. Goldman), Amer. Wildhfe D. Acad. Sci., 13:108-111. Inst., Washington, C, xiv-|-358 pp. WOOSTER, L. D. 1958. The bobcat of North America .... 1935. Notes on the effects of drought on Stackpole Co., Harrisburg, and Wild- animal populations in western Kansas. life Mgt. Inst., Washington, D. C, Trans. Kansas Acad. Sci., 38:351-352. 193 pp. AVAILABLE MISCELLANEOUS PUBLICATIONS IN MAMMALOGY UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY

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