UMl HARVARD UNIVERSITY LIBRARY OF THE Museum of Comparative Zoology JM 13 UNIVERSITY OF KANSAS miscellaneous MUSEUM OF NATURAL HISTORY ^^^^^^^™I^n No. 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 Reithrodontomys megalotis dychei J. A. Allen 90 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 Mustela frenota alleni ( Mcrriam ) 130 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 as a mountainous island surrounded by a sea of grass. The mesic climate, conif- erous forests, rugged and dissected to- pography, and diversity of geological structures and edaphic features in the Black Hills contrast sharply with the adjacent Northern Great Plains, which are characterized by semi-arid to arid grasslands and gently rolling topography. In some areas, general zones of transition tend to compromise the distinctiveness of these two physiographic entities. In spite of the northern and montane affinities of the Black Hills, the indigenous mamma- lian fauna is heterogeneous in origin. A definitive biogeographical analysis of the mammalian fauna has not been at- tempted prexiously. The principal purposes of this report are: 1 ) to delimit and describe the mam- malian fauna of the Black Hills of South Dakota and Wvoming (43 10'-44"50' N lat.; 103 20'- 104 50' W long.) as a nat- ural zoogeographic unit; 2) to describe the autecology and distributional pat- terns of each mammalian species in the Hills; 3) to discuss the geographic varia- tion and inferred speciation of these mammals; and 4) to analyze and inter- pret the probable biogeographic affinities of various species in light of proposed changes in late Pleistocene and Holocene environments. Thus, this study repre- sents a synthesis of systematic, zoogeo- graphic, ccologic, and historic factors and their bearing on the contemporary mammalian fauna of the Black Hills. The Recent Black Hills mammalian fauna comprises 62 species in 44 genera and six orders. Three of the species have been extirpated by man and subse- quently reintroduced to the Black Hills from other areas. Four other species (indicated by an asterisk in the Con- tents) are adventives; of these, two were introduced from North America and two from outside North America. Also in- cluded in the accounts are 25 species whose occurrence in the Black Hills is questionable or undocumented at pres- ent. ACKNOWLEDGMENTS Many persons have contributed di- rectly or indirectly to the completion of this paper. I am particularly grateful to the following persons for the loan of specimens, for information concerning specimens, or for permission to examine specimens: S. Anderson, K. F. Koopman, and R. G. \^an Gelder, American Mu- seum of Natural History; J. C. Moore, Field Museum of Natural History; R. R. Mansfield, Minnilusa Historical Museum; W. H. Burt and E. T. Hooper, Museum of Zoology, University of Michigan; J. D. Druecker and ]. S. Findley, Museum of Southwestern Biology, University of New Mexico; R. A. Martin, Department of Biology, South Dakota School of Mines and Technology; D. W. Block, E. J. Hugghins, and R. J. Walstrom, Department of Entomology and Zoology, South Dakota State University; B. Har- rell. Department of Biology, Uni\'ersity of South Dakota; C. O. Handley, Jr., D. A. Schlitter, and H. W. Setzer,' U. S. National Museum; R. H. Manville and J. Paradiso, U. S. Fish and Wildlife Serv- ice ( Biological Surveys Collection ) ; C. A. McLaughlin, Department of Zoology, University of Wyoming; and N. R. Whit- ney, Rapid City, South Dakota. I am indebted to Park Superintendent W. D. Hotclikiss and Chief Park Naturalist G. B. Robinson of Wind Cave National Park for pro\iding physical facilities and access to records on file at the Park. I wish to extend special recognition to J. A. King, Department of Zoology, Mich- igan State Uni\'ersity, for making a\ail- able his unpublished field notes and 6 TURNER: MAMMALS OF THE BLACK HILLS supplementary observations which great- ly enhanced my study. I also wish to thank R. Barbour and W. H. Daxis, De- partment of Zoology, Uni\ ersity of Ken- tucky, for assistance in the field in the summer of 1968. Some 46 persons associated with the Museum of Natural History of The Uni- versity of Kansas participated in field work in the Black Hills from 1947 to 1967. I want especially to recognize S. Anderson, R. H. Baker, E. L. Cockrum, E. R. Hall, J. K. Jones, Jr., C. A. Long, and H. W. Setzer, whose expeditions resulted in specimens and comprehen- sive field notes used in preparation of this report. For identification of ectoparasites, I am indebted to K. C. Emerson, U. S. Army; E. Garret, Bishop Museum, Hono- lulu; C. E. Hopla, Department of Zool- ogy, University of Oklahoma; G. M. Kohls, Rocky Mountain Laboratory, Hamilton, Montana; R. B. Loomis, De- partment of Biology, Long Beach ( Cali- fornia) State College; B. McDaniel, De- partment of Entomology and Zoology, South Dakota State University, Brook- ings, South Dakota; B. V. Peterson, Can- ada Department of Agriculture, Ottawa; R. D. Price, Department of Entomology, Fish, and Wildlife, University of Minne- sota; the late R. L. Usinger, Department of Entomology, University of California (Berkeley); N. Wilson, Department of Biology, University of Northern Iowa, Cedar Falls; and W J. Wrenn, Depart- ment of Entomology, The University of Kansas. The cooperation of the South Dakota Department of Game, Fish and Parks is fully appreciated. Scientific collecting permits were kindly issued by V. Johnson and J. W. Sprague. Chief of the Division of Game Management, J. Popowski, and State Game Biologists, L. Petersen and A. H. Richardson, contributed significant information concerning the artiodactyls and carnivores in the Black Hills region. State Game Wardens R. Butterfield, O. Meadows, C. Webster, and E. L. Woods also donated specimens and information pertaining to the larger fur-bearers in their respective districts. I am especially grateful to Assistant Chief Ranger J. F. Devenport, Wind Cave National Park, for contributing his firsthand knowledge of wildlife within the Park and adjacent regions. The advice of A. C. Meland, Soils and Water Conservation Service, concerning the edaphic characteristics of the Black Hills region is appreciated. A number of personnel in the U. S. Forest Service, Black Hills National Forest, ad- vanced my study in innumerable ways. Among these I would mention especially I. Case, W. D. Cloud, F. Fichtner, A. W. Jones, D. Kocer, G. Roby, K. C. Scholz, and J. C. Windsor. I am indebted to all those listed above and others unnamed, who gave generously of their time and efforts in my behalf. The National Science Foundation (through fellowships to student partici- pants for field work in June and July of 1965 and 1967 under Grant GE-7739), The Museum of Natural History, the Department of Zoology, and the Com- mittee of Systematics and Evolutionary Biology (NSF Grant GB-4446X) at The University of Kansas, and a Watkins Mu- seum of Natural History Grant, all helped to defray cost of field operations. A computer grant from the Department of Systematics and Ecology enhanced statistical analysis of the data. I am grateful to other friends and associates, especially Elmer C. Birney, John B. Bowles, Hugh H. Genoways, and Carleton J. Phillips, who have given of their time and criticisms, and I am obliged to Barry L. Siler for preparing the illustrations. To my wife, Barbara M. Turner, I give special recognition for her patience and perse\'erance during the final phases of the project and for typ- ing the first draft of the manuscript. Similar recognition is given to Mary H. Michener for reading the final proof. I am indebted to Robert S. Hoffmann and Ronald McGregor who supported my investigation through encouragement and contribution of crucial counsel. In addition, Wakefield Dort, Jr., and A. W. MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY Kuchler read and criticised portions of the manuscript. I extend my thanks to Howard J. Stains, Southern IlHnois Uni- versity, who first instilled in me an in- terest in zoology, and through his teach- ing and personal example gave great impetus to my formal education. Especially instrumental in the forma- tion and completion of my study was J. Knox Jones, Jr., who gave continued guidance throughout the duration of the project, devoted many arduous hours to counseling, assisted in field problems, and aided in preparation of the com- pleted manuscript. His unselfish assist- ance is acknowledged with sincere grati- tude. Professor Jones pro\'cd himself to be capable of considerable patience, the depths of which only I can truly appreci- ate. HISTORICAL resume: Little is known about prehistoric man in the Black Hills area. Archeological findings from several sites indicate that early inhabitants of the area (7000-9000 BP) were bison hunters organized so- cially into small hunting bands, each composed of several cooperating families (Black Hills Area Resource Study, 1967: 21). Several Indian tribes (in chrono- logical sequence: the Poncas, Kiowas, Crow, and Cheyennes) lived near the Hills at different times. When white men first came, the region was in the posses- sion of the Teton- Dakotas, more com- monly called the Sioux. However, the Sioux were immigrants, drifting west from the Great Lakes country and dis- placing the Cheyennes as late as the mid-eighteenth century. The Black Hills region is rich in history, much of which has been re- corded in accounts of the pioneers that settled there. I have not attempted to consult the vast historical literature, but rather have restricted myself to accounts of the various scientific and military ex- plorations of the study area. Results of these expeditions are recorded in several Reports of the Secretary of War in the Executive Documents of Congress. Ad- ditional information can be obtained from the collections and publications of the South Dakota State Historical So- ciety, Pierre, South Dakota 57501. Francois and Louis-Joseph Veren- drye, two French explorers, entered the Black Hills in early February of 1743; they were probably the first white men to penetrate the region. Although Meri- wether Lewis and William Clark did not enter the Hills proper, a French trader, Valle, encountered the explorers near the mouth of the Cheyenne River and told them of the "Black Mountains" to the west. Having received similar re- ports while camped at the mouth of the Bad River in 1833, Prince Maximillian termed this pine-clad range the "Black Hills" (Thwaites, 1906). He also in- cluded the Killdeer Mountains and Little Missouri River Badlands of North Da- kota under this designation ( Bailey, 1927:25). Both Hunt's American Fur Company Astoria Expedition (1811) and Jedediah Smith's expedition of 15 fur traders (1823), passed through parts of the Black Hills. Additionally, there was a number of other traders and trappers who traxersed the Hills during the first half of the nineteenth century, but these men left little record of their passing. Fur trading posts were established near the mouth of the Belle Fourche Ri\'er and on the White River, near the mouth of Wounded Knee Creek, as early as 1828. Scientific exploration of the region began in 1852, when Dr. John Evans mapped the Badlands and the eastern foothills under the auspices of the Da\'id Dale Owens Geological Survey. In March 1853, Congress appropriated funds for a survey of xarious proposed routes, along \\'hich a railroad might be constructed from the Mississippi River to the Pacific Ocean. Survey parties were organized by the War Department, and supplies for collection of natural history objects were pro\'ided by the Smith- sonian Institution. The first known speci- mens of mammals from the Hills were TURNER: MAMMALS OF THE BLACK HILLS taken b\ these expeditions. Topographi- cal and geographical knowledge of the country along thc> White and Bad rivers was incremented in 1855 with the mili- tary expedition headed by General Wil- Ham S. Harney, and including topograph- ical engineer Lieutenant G. K. Warren, and geologist Dr. F. V. Hayden. This party passed along the southern periph- ery of the Black Hills from Fort Laramie enroute to Fort Pierre (Warren, 1856). Two years later, Hayden again accom- panied Lt. Warren to the region, record- ing observations on 44 kinds of mam- mals (Hayden, 1859). Entering from the south by way of Stockade Beaver Creek (Weston County), they traveled along the western edge of the Hills to Inyan Kara Mountain, Crook County, Wyoming (Fig. 2). Here they were turned back by threatening bands of Hunkpapa and Miniconjou Sioux; they tra\'eled southeastward to the vicinity of Rapid City, then north to Sturgis and Bear Butte and returned to Missouri via the Niobrara River (Warren, 1856; 1859; Hayden, 1856; 1859). Hayden (1862: 138-151) summarized his observations on the natural history of the Upper Mis- souri based on excursions from 1854 to 1862 commenting on 59 kinds of mam- mals, including representatives from the Black Hills. Some reports of travel through the Black Hills in the mid-nineteenth century must be viewed with skepticism, because in those days the term "Black Hills" also applied to the Laramie Range along the North Platte River in Wyoming. For example, in 1856, Lieutenant F. T. Bryan and naturalist W. S. Wood were part of an expedition that was instructed to build a road from Fort Riley, Kansas, to Bridger's Pass, Wyoming (Bryan, 1858). Although they did not enter the Black Hills, some of the specimens of mammals collected by Wood bear this locality on the labels. Many books and articles have been published concerning the Oregon Trail of Francis Parkman in 1846; the numerous references to the Black Hills in these publications actually are appli- cable to the Laramie Mountains (Wade, 1947:393). Hayden, with Captain W. F. Ray- nolds of the Yellowstone Expedition, passed through the northern Black Hills in 1859, reaching Devil's Tower on 20 July ( Raynolds, 1868 ) . Both the eastern and western margins of the Hills were reconnoitered by elements of the Pow- der River Expedition of 1865. In spite of the pressure to open up the Hills for exploitation of gold, the Indian Treaty of the Peace Council of Laramie, signed on 29 April 1868, gave that part of the Dakota Territory lying west of the Mis- souri River to the Sioux, temporarily clos- ing the frontier. Increasing rumors of gold in the Black Hills and continuing troubles with the Indians resulted in the expedition of Brevet Major General George A. Custer in 1874 (O'Hara, 1929; Jackson, 1966). Members of this expedition traveled about 600 miles in 60 days, and returned with numerous photographs, many ob- servations on the natural history of the region, and gold from French Creek, Custer County. Dr. Othniel C. Marsh, the noted paleontologist, was invited to accompany Custer, but instead sent along his young assistant, George Bird Grin- nell, who took notes on 34 species of mammals (Grinnell, 1875:79-84). The expedition entered the Hills from the north, traveling down Castle Creek, south to the Cheyenne River. It re- turned northward to the area around Harvey Peak, camping at French Creek near the present town of Custer, and then left the Black Hills by way of Box- elder Creek and Bear Butte. Ludlow ( 1875 ) gave an excellent report of the reconnaisance, and included two foldout maps of the routes of Warren in 1857, Raynolds in 1858, and Custer in 1874. The Newton-Jenney U. S. Geological Sur- vey of 1875, with Colonel Richard I. Dodge commanding the military escort, spent five months in the Hills studying the geology and natural resources of the area (Jenney, 1868; Newton and Jenney, 1880). Due to ill health, C. G. New- 10 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY berry, the naturalist assigned to the ex- pedition, was compelled to resign at Fort Laramie on the eve of departure; thus the only records of mammals ( 15 kinds ) observed or taken on the trip were those recorded bv Colonel Dodge (1876: 120- 123, 128-i;34). As the Black Hills opened to white settlement, biological exploration intensi- fied. Wetmore (1968:215-216), and Pet- tingill and Whitney (1965:16-17) have reviewed botanical and ornithological in- vestigations, respectively. In the sum- mer of 1894, Walter W. Granger left the paleontological expedition sponsored by the American Museum of Natural His- tory to the Badlands of South Dakota. He journeyed west to the Black Hills and collected 22 species of mammals, mostly from the southeastern and central sections (J. A. Allen, 1895a), including several undescribed kinds (J. A. Allen, 1894a, 1894b, 1895b). From 1899 to 1911, Henry Behrens made a collection of 30 kinds of mammals on his ranch along Spring Creek, and along the foot- hills south of Rapid City. Fifty-five of these specimens now are housed in the Pioneer Room of the Minnilusa Histori- cal Museum in Rapid City, South Da- kota. The U. S. Biological Survey became active in the region at the turn of the century (Merriam, 1888, 1889, 1891). Vernon Bailey and Merritt Gary were re- sponsible for obtaining mammals and natural historv data from the region. Be- tween 1899 and 1912, Cary (1917) peri- odically investigated the area along the South Dakota-Wyoming border in the vicinity of Elk Mountain. Bailey (1914) intermittently collected in the Black Hills from 1887 until 1913; observations of 25 species of mammals were made on his initial excursion near Rapid City and Deadwood late in 1887 (Bailey, 1888). The materials, gathered by Granger, Bailey and Gary contributed significantly to the present study. A. H. Howell, N. Dearborn, and a field party from the U. S. National Mu- seum worked in the central part of the Hills from mid-May to mid-June of 1910, and P. Moulthrop and G. W. Phillips from the Cleveland Museum of Natural History collected in the same area in August 1929 (Bole, 1935; Moulthrop, 1936). Victor H. Cahalane (1948, 1951), as Acting Chief of the Wildlife Division at Wind Cave National Park, compiled a partial list of mammals of that area in the course of field work from 15 August 1935 to 10 February 1936 (on file at Wind Cave National Park). Somewhat later, A. M. Stebler (1939) and L. R. Dice ( 1939 ) from the University of Michigan Museum of Zoology, initiated field investigations that resulted in pub- lished reports on mammals from the Black Hills. J. A. King (1951, 1955), also from the University of Michigan, worked in the central part of the Hills and at Wind Cave National Park at intervals from 1945 to 1952, and C. B. Koford ( 1958 ) , from the Museum of Vertebrate Zoology, University of California, Berke- ley, carried out investigations in the Wyoming sector, principally in the \ icin- ity of Devils Tower. In 1947, 1951, 1961, 1965, and again in 1967, field parties from the Museum of Natural History of The University of Kansas collected mammals in western South Dakota and northeastern Wyo- ming (Jones and Packard, 1958; Long, 1965; Jones and Genoways, 1967a, 1967b; Turner and Jones, 1968; Turner and Davis, 1970; White, 1952, 1953a, 1953b). My own work in the Black Hills began in the summer of 1965; I returned to the study area throughout the summers of 1967 and 1968. A week was spent in quest of hibernating bats in late Novem- ber of 1967, and a week each in obtain- ing photographs and samples of soil in August of 1969, and March of 1970. In all, I was in the field for 31 weeks from mid-June 1965 to March 1970, including two months as a Ranger-Naturalist at Wind Gave National Park in 1968. Ad- ditionally, tvvo weeks were required in 1968 to examine specimens from the Black Hills housed in museums other than at Kansas. TURNER: xMAMMALS OF THE BLACK HILLS 11 METHODS AND MATERIALS Each of the six orders of Black Hills mammals and the 19 families ( discussed briefly), including 44 genera and 62 species, are arranged in text following the arrangement of Hall and Kelson (1959). The species of each genus and, where appropriate, the subspecies of each species, are in alphabetic order. Keys to species, disti-ibution maps, and enumeration of characters applicable to the \arious ta.xonomic categories, are not pro\ ided herein as these were deemed to be readily accessible in other publi- cations. The account of each monotypic spe- cies or subspecies native to the Black Hills incorporates the following: 1. The scientific name, employed in agreement with the International Rules of Zoological Nomenclature, and fol- lowed on the same line by the name of the author. 2. The vernacular name, in general accord with Hall ( 1965 ) , and that which is considered appropriate for all sub- species of the given species. 3. The synonymy, in which the first citation is to the original description and is followed by designation of the type locality. The second citation is to the first usage of the currently accepted name-combination employed herein, un- less that combination is identical to the name as originally proposed. In a few cases, a third citation is to a taxon de- scribed from the Black Hills, but now placed in synonymy. 4. The disirihuiion, which concerns the regional distribution of each species and comments pertaining to the abun- dance and apportionment of each throughout the Black Hills. Included here are notations on ecological, altitudi- nal, and seasonal distributions; the latter is especially important for those mam- mals that hibernate or migrate. 5. Comments on systematics are in- cluded in most accounts. Comparisons of characters are made between the Black Hills populations and those of sur- rounding areas when these are warranted for taxonomic clarification or when con- spicuous geographic variation is evident. 6. All measurements, external and cranial, are given in millimeters. Stand- ard external measurements were read from labels attached to the specimens, excepting length of forearm; the latter is applicable only to bats and was meas- ured from prepared museum specimens. Weights of adult males and non-pregnant adult females are given in grams. Stated measurements are the arithmetic mean, followed by the standard deviation. Un- less otherwise stated, measurements are of adults only. Cranial measurements were taken to the nearest tenth of a millimeter with dial calipers in the man- ner described by Hall (1946:672-685), and Packard (1960:584-585); identical measurements were not taken for all species. Variation in color was assessed by direct comparison of specimens and also by use of a Photo\'olt Photoelectric Re- flection Meter, Model 610, utilizing red, green, and blue filters. Readings were taken from the middorsal region of mu- seum study skins and could be repeated on the same individual with minimal variation. Samples of soil from various collecting sites were first dried for sev- eral hours in an oven, then subjected to color analysis with the reflection meter. Resultant measurements of reflected light were recorded for each skin and soil sample as percentage values of pure white calibrated against a standardized block of magnesium carbonate. These recorded measurements actually repre- sent a composite value of both intensity of hue (that is, more red or less red in color) and tone (paler or darker in color ) . For example, pale brownish soils yielded higher reflectance readings when using a red filter than did dark reddish soils. Reflectance readings obtained from all three filters were summed in order to achieve an approximate index of tone. If, for instance, the specimens or soil samples were arranged in order of de- creasing paleness of overall coloration, the resultant sequence derived by direct 12 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY comparison would be in agreement with the summed total (trichromatic) reflect- ance, or tone (Figs. 13 and 14). Inten- sity of each hue was derived by dividing the respective initial reflectance reading from each filter by the total reflectance readings from all three filters. Extremely greasy or damaged skins were not in- cluded in the color analysis. In analysis of geographic variation, specimens were segregated by age, sex, season, and geogi'aphic origin. The measurements obtained under these groupings were then subjected to stand- ard univariate statistical analysis and to an overall Analysis of Variance. If the Analysis of Variance indicated that sig- nificant differences existed among groups, then the Sums of Squares Simul- taneous Test Procedure of Gabriel (1964) was applied in order to locate these differences. 7. Remarks on autecoIog,i/ ordinarily include information pertinent to the nat- ural history of each species. Methods of capture, time of activity (i.e.. Mountain Daylight Time, MDT, or Mountain Standard Time, MST), description of habitat, enumeration of associates, and additional noteworthy observations are recorded. A brief chronicle is given for those species that were common in his- toric times, but now have been reduced in numbers or extirpated. Reproductive data are summarized for each species for which information (usually taken from specimen labels or field notes of the collector) is available. Molt data were obtained from examination of museum specimens, both diy and in alcohol, by directing a stream of air through the pelage and noting the presence or ab- sence of underlying new hairs. In recent year, specimens were examined in the field immediately after capture for ecto- parasites. Parasites collected in this manner were preserved in 70 percent alcohol and referred to various special- ists for identification. In compiling the above-mentioned data and observations, field notes of 46 individuals were con- sulted, in addition to my own firsthand observations. 8. The records of occurrence, include both specimens examined (based on 4727 specimens) and additional records. Under specimens exatnined, the first notation is the total number of speci- mens examined by me, followed by the exact locality of capture, the number of specimens from each locality, and desig- nation of site of specimen deposition. Localities are allocated to their respec- tive counties, which are grouped under either South Dakota or Wyoming. County names and the localities within each county are arranged from north to south (and \\est to east if more than one locality occurs in the same latitude within a county). The many specimens examined for comparisons from areas surrounding the Black Hills are not enu- merated. Abbreviations designating specimens examined in collections other than the Museum of Natural History of The University of Kansas are as follows: AMNH — American Museum of Natural History, New York; FMNH— Field Mu- seum of Natural History, Chicago; MHM — The Pioneer Room, Minnilusa Histori- cal Museum (collection of Henry Beh- Rapid City; MSB — Museum of rens Southwestern Biology, University of New Mexico, Albuquerque; NRW — Col- lection of N. R. Whitney, Rapid City; SDMT— Department of Biology, South Dakota School of Mines and Technology, Rapid City; SDSU— Department of En- tomology and Zoology, South Dakota State University, Brookings; UK — De- partment of Zoology, University of Ken- tucky, Lexington; UMMZ — Museum of Zoology, University of Michigan (in- cluding specimens formerly housed in the Cleveland Museum of Natural His- tory), Ann Arbor; USD — Department of Biology, University of South Dakota, Vermillion; USNM— United States Na- tional Museum (including collections of the U. S. Biological Surveys), Washing- ton, D. C; UW — Department of Zoology, Uni\'ersity of Wyoming, Laramie; WCNP— Collection of Wind Cave Na- tional Park, Hot Springs. Additional records consist of reports TURNER: MAMMALS OF THE BLACK HILLS 13 from the literature, specimens in collec- tions that I have not examined, or identi- fications of presumed accuracy as re- corded in card files of the U. S. Biologi- cal Sur\'cy (USES); the abbre\'iation BB designates specimens in the collection of B. Bailey that were identified by per- sonnel of the Survey. Localities from which specimens were examined are not duplicated in the additional records. Most sites from which mammals were obtained are shown in figure 2. ENVIRONMENT GEOGRAPHY The Black Hills constitute a maturely dissected, isolated, mountainous region of approximateh' 4000 square miles that resulted from intermittent domal uplifts in Cretaceous, Miocene, and Pleistocene times. The area extends 120 miles in a northwest-southeast direction, and is 50 miles wide at the widest point. The Hills are entirely surrounded by the non-glaci- ated Missouri Plateau section of the Northern Great Plains physiographic province (Fenneman, 1931), and rise abo\e the plains to an elevation of about 4000 feet on the east and 3000 feet on the west. The highest point (Harney Peak) lies 7242 feet above sea level. Most of the region lies within the drain- age of the Cheyenne River, which circles the south end of the Black Hills, and the Belle Fourche Ri\'er, which skirts the north edge of the area (Figs. 1 and 2). As defined herein, the Black Hills are delimited by the distribution of Jurassic shale and sandstone of the Sundance Formation ( Fig. 3 ) . Thus, the region includes the Black Hills proper, as well as the Bear Lodge Mountains and Devils Tower of Wyoming; the latter two areas are closely allied with the Hills. Ranges of mountains nearest the Black Hills are the Laramie Mountains (elevations to 10,272 ft) and Big Horn Mountains (ele- vations to 13,165 ft); these lie approxi- mately 150 miles to the southwest and west, respectively, in Wyoming. GEOLOGY AND PHYSIOGRAPHY As a result of the presence of a di- versity of geological structures and pro- ductive mineral deposits, there are nu- merous publications concerning the geol- ogy of the Black Hills. The reader is directed to Darton (1909), Darton and Paige (1925), Mcintosh (1931), Tullis (1951), and Cries and TulHs (1955) for detailed geological accounts of the re- gion. The Black Hills uplift is a crescent- shaped, anticlinal dome that rises sev- eral thousand feet above the surrounding Northern Great Plains (Figs. 1 and 3). The Central Basin is located slightly east of the main axis of the dome. It is a rugged mountainous core of Pre-Cam- brian igneous rock and Pre-Cambrian, Paleozoic and Mesozoic sediments. The Basin is interspersed with park-like val- leys and steep, narrow canyons that in- cline to the north, south, and east. In the Harney Peak-Needles-Mount Rushmore area ( Fig. 4 ) there are bare granitic and mctamorphic ridges. Encircling the Cen- tral Basin is the Limestone Plateau of rolling highlands (Figs. 5 and 10). The plateau is about two miles in width to the east, and reaches a maximum of 15 miles in width in the northwestern por- tion of the Black Hills. It is comprised of Paleozoic sediments (sandstone, shale and limestone ) of the Deadwood, White- wood, Pahasapa-Englcwood and Minne- lusa formations. The wall-like escarp- ment of the plateau reaches elevations of 7100 feet; it faces inward toward the Central Basin and is interrupted by such stream valleys as Sand, Cold Spring, and Spearfish creeks. In some areas (e.g., Big and Little Spearfish Canyons; Fig. 6) of the Limestone Plateau, high cliffs pro- ject above the valley floors. The rugged terrain in the northern Central Basin and northwestern Limestone Plateau was formed by solidification of igneous in- 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 are broadly rolling tablelands that com- prise the foothills and the Foothill Tran- sition Soil Association ( see SOILS section below). The foothills are less extensive along the eastern slope due to a steeper gradient, and are composed of Paleozoic limestone (Minnekahta Formation) and TURNER: MAMMALS OF THE BLACK HILLS 15 104' NEWCASTLE f NIOBRARA CO. 20 Hl MILES _ PIACERVIILE PACIOLA '/ \_^ C_N_^_ "" — a hori- zon that is weakly platy in structure and from four to 12 inches in depth. 4) Bi — a four-inch-thick transitional layer between the Ao and B2 horizons; Bi is absent in the Hills. 5) B- — a horizon which is brown and blocky in structure and pos- sesses clayfilms on all surfaces; this layer is between 10 and 30 inches in depth. 6) C — a horizon which varies in compo- sition in different areas. Other zonal soils that occur on lower sloDes are the Chernozems and Chestnut soils. Color of the surface layer of the Chernozems is black to dark, grayish brown. Soils of this type occupy open woodlands and are paler in color and browner in the southern part of the Hills than in the northern sections. Ai hori- zons usually are five to eight inches in depth and granular in structure. Bo horizons arc about eight to 10 inches in depth and structured of prisms that ordi- narily are coated with clayfilms on all surfaces. B.^ horizons have a weak, pris- matic structure; they are usually more than six inches in depth and more clayey in composition. Both the B... and C hori- zons are enriched with lime leached from the upper layers. The surface layer of Chestnut soil is brown or reddish in color and developed under grasslands. The A, horizon is organic and about two to four inches in depth. The B- horizon is be- tween 10 and 15 inches in depth and of prismatic structure with clayfilms on most surfaces. The B.-; horizon is at least six inches in depth and structured of prisms. Both the B.. and C horizons contain free carbonates. Azonal soil groups in the Black Hills include Regosols, Lithosols, and Allu- vium. Regosols are thin soils developed in unconsolidated material, whereas Lithosols are thin soils over solid rock (within 1(S inches of the surface). These immature soils occupy steep slopes where runoff is so excessive that little leaching or development of humus occurs. No true subsoil is present. Alknium occu- pies flood plains of various water courses in the Hills. Alluvium tends to be darker and more humic than soil of the adjacent uplands and frequently supports a not- ably different riparian plant community. A great need exists for detailed infor- mation on soils in the Black Hills. Most of the soils are unclassified and are stony or rocky, reflecting the local geology and physiography. For detailed information concerning land use of the soil associa- tions, the reader is directed to Austin (1965:26-28), Westin ef al. (1967:1-32) and to miscellaneous publications issued by the Soil and Water Conservation Dis- tricts of the respective counties. Some relationships of soils to the distribution and speciation of mammals in the Black Hills are discussed bevond. The following description of Soil As- sociations in the Black Hills (referred to in text) has been synthesized from frag- mentary literature and in consultation with the South Dakota Soils and Water 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 Description, classification, and utiliza- tion of lakes and streams in the Black Hills have been discussed in the Black Hills Area Resource Study of 1967, and by Stewart and Thilenius (1964). All water courses that arise in the Black Hills comprise 1302 miles of sh-eam drainage, and empty either into the Belle Fourche or Cheyenne rivers; both rivers originate on the gently rolling plains to the west in Wyoming. The Belle Fourche River, with a surface drainage of 7210 square miles, borders the northern mar- gin of the Hills; whereas, the Cheyenne River, with a surface drainage of 2000 square miles, flows along the southern periphery. Together, these rivers have an annual flow of 522,000 acre-feet; they join on the prairie about 60 miles east of the Black Hills and flow eastward (as the Cheyenne) into the Missouri River. The northwest-southeast alignment of their major tributaries, such as Boxelder, Rapid, Spring, Battle, French, and Beaver creeks and of other streams in western South Dakota, coincides with the prevailing wind direction, and ap- pears to be the result of periodic accumu- lation of locally derived eolian sediments 26 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY (White, 1961:207). Although June is the time of maximum precipitation, max- imum surface runoff is in May owing to the greater evapotranspiration rates en- countered later in the growing season. Floods are common in both months. Due to abrupt topographic relief, most streams in the Black Hills are radial-dendritic systems that flow swiftly; only in a few montane meadows and broad valleys does a more gradual gra- dient allow some meandering and for- mation of small, semimarsh areas ad- jacent to stream channels. In addition to aquatic mammals such as the beaver (Castor canadensis) and muskrat (On- datra zihethicus), riparian species such as the mink (Mustela vison), masked shrew, jumping mouse, long-tailed vole, and meadow vole inhabit these moist environs. When European man first entered the Black Hills, beaver ponds were the only sources of standing water. Now the foothills are interrupted by thousands of small reservoirs that store water for live- stock and for fish and wildlife; major impoundments such as Angostura, Deer- field, Pactola, Keyhole, and Belle Fourche reservoirs also have been con- structed. These artificial bodies of standing water (usually located over shist or shale) are used for recreation, as well as for watershed development and irrigation. Except for the headwaters, all streams in the Black Hills are affected to some degree by pollution. Whitewood Creek is the most severely polluted stream in the Hills; it carries an extremely high content of rock-flour and chemical pollu- tants from mining operations and a heavy load of municipal wastes from Lead and Deadwood. The North Fork of Rapid Creek also is polluted with a high con- tent of bog-iron wastes. Stockade Lake is of poor equality due to sewage effluents from Custer. A dense bloom of a toxic blue-green algae (Aphanizomenon flos- aqiiae) results from the supplemental nitrogen and phosphorous, and drasti- cally reduces productivity of the lake (Stewart and Thilenius, 1964:46). Dodge (1876) recorded the hydrog- raphy of the Black Hills when he ac- companied the Newton-Janney U. S. Geological Survey of 1875. Stream flow was reported as substantial and the wa- ters as being "cool, clear, and pure." Al- though riparian vegetation was suffi- ciently dense to impede travel along the stream bottoms, forest cover evidently was less extensive then than now, in that large older trees were "scarcely to be found." Adequate fire protection and management practices such as "thinning" have allowed a mature forest with a closed canopy to develop over much of the Hills. Thickening of the forest, with resultant interception of precipitation and increased evapotranspiration, is the greatest single factor in reduction of total surface discharge and in decrease of moisture available for recharging ground water stores (Stewart and Thil- enius, 1964). Trampling of riparian vegetation by cattle around springs and creeks, construction of roads along stream channels, tapping of ground wa- ter supplies with wells, storage and utili- zation of water for agriculture and indus- try, and deposition of tailing from mines, tannic acid from saw mills, and wastes from municipalities continue to degrade aquatic habitats in the Black Hills. VEGETATION The most conspicuous component of the Black Hills vegetation is the conif- erous forest, which is dominated by a montane belt of ponderosa pine (Pintis ponderosa) (Fig. 9). In more mesic sites on northern exposures in the Central Ba- sin and on the Limestone Plateau, there is a subalpine belt of white spruce ( Ficea iijauca) with an associated understory of plants with northern affinities (Fig. iO). Kentucky bluegrass (Poa prafensis), in- troduced from Europe in the 17th Cen- tury, is the predominant graminoid plant of the open meadows, parklike areas, and grasslands of the foothills and Red Val- ley. The flora is a mixture of boreal. TURNER: MAMMALS OF THE BLACK HILLS 27 cordillrran, eastern deciduous forest, and Great Plains species (Rydberg, 1896; Webb, 1965). Biogeographic affinities of the Black Hills biota ha\e been treated hv \arious authors. Birds are discussed bv Miller (1941), Pettingill and Whit- ney (1965), and Mengel (1970). Byers (1961) and Ross (1965; 1970) dealt with insects, and Smith ( 1957 ) discussed the smooth green snake. Willo\\'s and lichens were treated by Froiland ( 1962 ) and \\'etmore (1968), respectively. Forest and other vegetational components are discussed by Buttrick ( 1914 ) , Halliday and Brown (1943), Havward (1928), Mcintosh (1931), Dillon' (1956), Potter and Green (1964), and Watts and Wright (1966). The most exposed sections in the Black Hills, such as the crown of Harney Peak and the Needles section, either are de\'oid of ^'egetation or support bryoids (mosses) and thallophytes (lichens). A climatic timberline is not present; in- stead, the absence of trees along the high summits is due to lack of soil and to extreme exposure. In general, three vegetational types occur in the Black Hills — coniferous forest, deciduous woodlands, and prairie grasslands. Dry slopes are dominated by ponderosa pine, the dark appearance of which accounts for the Sioux name PaJia Sapa or "Black Hills." Usually broadleafed trees are de- veloped mainly along drainage channels, or are present as groves on old, burned areas. Grasslands are found mostly on slopes of the foothills, but also in isolated prairie areas even at higher elevations. Coniferous Forest Association. — This association forms a forest that is domi- nated by ponderosa pine, which appears to be extremely tolerant of xerophytic conditions (Fig. 9). The species ranges between elevations of approximately 3500 and 7000 feet, and forms parklike forests at higher elevations that give way to open woodlands at lower elevations. On the pine-clad uplands, the soil is coarse, well drained, and quite warm in the summer. In the western section, and especially on the hogback, western red cedar (Jiiniperus virginiana) intrudes among the pine. Along the cooler canyon floors and northern slopes of the central and moist northern sections of the Black Hills, the coniferous forest includes white spruce, paper birch (Betida papyifera), and quaking aspen (Populus tremu- Joides) . The accompanying understory consists of wild sarsaparilla (Aralia nti- (Ucoidis), twin-flower (Linnaea hore- cdis), swamp currant (Rd)es Jactistre), bunchberry (Cornus canadetisis) , red osier (C. stolonifera) , Venus' slipper (Calypso hidhosa), squashberry (Vi- hurnum edide) buffaloberry (Shep- Jierdia argentea), and huckleberry (Vac- cinium memhranaceum) in close asso- ciation. White spruce may occasionally dominate the coniferous forest of lower slopes in these areas, but usually it oc- curs in subalpine belts that are in close proximity to montane belts of ponderosa pine (Fig. 10). Predominant associates of the pine are various bunch grasses, bearberry [ArctostaphyJos uva-ursi), wild rose (Rosa acictdaris), creeping juniper (Juniperus horizontalis) , ground juniper (/. communis), Oregon grape (Berlyeris repens), redroot (Ceanothus velutintis) , New Jersey tea (C. ovatus), and poison ivy ( Rhus radicans ) . Deciduous Woodland Association. — Bur oak (Que reus macrocarpa) com- monl\' is intermixed with the pines along drainages near the periphery of the Hills; it occurs in drier valleys, on drier slopes, and on sandy to gravelly soils. Quaking aspen, paper birch, green ash (Fraxinus pennsylvanica) , and American elm (Ul- nius americana) are also present. In the northern section, and especially in the Bear Lodge Mountains, bur oak is the dominant tree growth on many slopes leading away from stream channels. The oak has an undergrowth of hawthorn (Crataegus chrysocarpa) , chokecherry (Pyrus virginiana), hop hornbeam (Os- trya virginiana) , and poison ivy. Riparian Association. — Streamside habitats in the Black Hills are character- ized by fluviatile soils that are cool, moist, and somewhat more finely tex- 28 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY tured than that of the pine-clad uplands. These habitats support American elm, cottonwoods (Poftiliis cleltoides), box- elder (Acer negundo), willows (Salix sp.), serviceberry (AmelancJiier aJnifo- lia), red-berried elder {?>amhucus pu- hens), blue flag [Iris missoiiriensis), Solomon's seal {Fohjgonaium hijlorum), carices {Carex sp.), and rushes (Junciis sp.). Mountain Prairie Association. — On a few rolling uplands at fairly high ele\'a- tions, an isolated prairie type of habitat develops in areas where soil texture, moisture, and depth reach a favorable balance. Examples of these grasslands are the Gillette and Reynolds prairies, and the Bald Hills (Fig. 11). Green needlegrass (Stipa viridiila), western wheatgrass (Agropyroti sinithii), blue grama (Boufelonra gracilis), timothy (Pldetim prafense), red-top grass (Ag- rostis palustris) and brome grass {Bro- mus sp.) occur with silver sagebrush {Artemisia cona), brittle pricklypear cac- tus (Opiintia fragilis), common prickly- pear cactus ( O. compressa) , plains prick- lypear cactus (O. pohjcantha) , pincush- ion cactus {Coryphantha missouriensis) , goldcnrods (Solidago sp.), Indian paint- brush (Castilleja sidpluirea), mariposa lily [CaJchortus nuttaUii), gland stem (Adenocaidon hicoJor), and asters {Aster sp.). Great Plains Prairie Association. — Many tongues of the surrounding plains extend into the Hills, especially from the south. The prairie-forest border is be- tween 3500 and 4000 feet in elevation. Graves (1S99) indicated that in areas where trees occur in xalleys and on north-facing slopes, the prairie-forest border is natural. However, at localities where trees occur on ridges instead of valleys, the prairie-forest border prob- ably is due to fire. Prairie grasslands formerly pre\'ailed in the Red Valley. Vegetation of the prairie is far from uniform in its composition. Dominants among the grasses are western wheat- grass, green needlegrass, blue grama, sideoats grama {Bouteloura ciirtipend- ula), little bluestem {Anclropogon sco- parius), big bluestem (A. gerardi), needle and thread grass {Stipa comata), porcupine grass (S. spartea), buffalo grass {Biichloe dactyloides), Indian grass {Sorga.struni avcnacciim), and Kentucky bluegrass. Forbs include pricklypear cactus, pin-cushion cactus, soapweed (Yucca glatica), sweet clover {Melilotus officinalis), blue vervain {Vervena has- tata), lamb's-quarters {CJienopodium al- bum ) , sunflowers ( Helianthus sp. ) , hare- bell (Campanula rotundifolia), western salsify (Trogonpogon dubius), prairie sandreed (Calamovilfa longifolia), skunkweed (Croton texensis), scarlet gaura (Gaura coccinea), and wild alfalfa (Psoralea tenuifiora) . Dominant browse species are ehokecherry, skunkbush su- ndae (Rhus trilobata), wormwood sage- brush (Artemisia dracuncidus and A. dracunuloides) , fringe sagebrush ( A. frigida), mountain mahogany (Cerco- carpus montanus), western sandcherry (Prunus besseyi), and wild rose (Rosa ivoodsii). Scientific names of plants in the resume follow the usage of Wetmore (1968) and Pettingill and Whitney ( 1965 ) , \\'hereas vernacular names gen- erally follow the usage of Over (1932). Professor Ronald L. McGregor and asso- ciates at The Universitv of Kansas cur- rently are studying the flora of the Black Hills region, and I am grateful to them for \erifying the nomenclature or sug- gesting appropriate changes for use here. PLEISTOCENE HISTORY Pre-Wisconsin Events. — The Pleisto- cene Epoch was characterized by a series of climatic fluctuations throughout the world. (At least four southern cool plu- vial periods occurred conciurently \\'ith northerly continental glaciation. Subse- quent to each of these episodes, a com- paratively warm, dry interglacial inter\'al followed.) As a consequence of these oscillations, distribution and speciation of many elements of the boreal and tem- perate biotas were markedly affected. Prior to the Pleistocene, mountain- TURNER: MAMMALS OF THE BLACK HILLS 29 building in the Rock\' Mountain region probably created a rain shadow in the Black Hills region. Further uplift during the Pleistocene caused entrenchment of streams around the edge of the Hills (Darton, 1909). Previously, the Chey- enne Ri\er and other streams to the south of the Hills probably flowed to- ward the Gulf of Mexico. Thus, the present course of the Missouri Ri\ er had not yet been attained (Flint, 1955; Lemke et al., 1965 ) . Because of relatively low elevation and low precipitation, montane glaciers did not form on the Black Hills (Hay- ward, 192S; Mcintosh, 1931). Nonethe- less, Darton (1906) and others reported that Cordilleran glaciers formed in the Bighorn Mountains between 9500 and 11,000 feet and flowed down to 6500 feet. There is no evidence that conti- nental glaciers entered Wyoming ( Long, 1965), but at its maximum, about 18,000 BP, the Wisconsin ice sheet terminated about 150 miles east of the Black Hills (see below), and that part of South Dakota west of the Missouri River es- caped direct glacial action ( Flint, 1957 ) . Intense frost evidently occurred within 50 to 100 miles of the ice sheet, making the Hills region subject to periglacial in- fluences. Erosion, deepening of stream channels, deposition of sands and gravels, and development of soils also occurred in that region during the late Pleistocene. The Nebraskan and Kansan glacia- tions covered most of South Dakota east of the Coteau du Missouri, whereas the Illinoian glacier reached only the ex- treme southeastern corner of the state, near Hartford (Lemke et al, 1965). This record is based principally on till found overlying presumed Pearlette Volcanic Ash of late Kansas or Yarmouthian age (Schultz and Smith, 1965). Distribution of Pre-Wisconsin till indicates that gla- ciers advanced chiefly via the James River and Red River lowlands. The Wis- consin ice front, as evidenced by accumu- lation of drift deposits, extended from southeastern South Dakota, northwest- ward toward North Dakota, just west of the Missouri River, roughly paralleling the river channel (Flint, 1957). Apparently, these glacial periods were characterized by cool, moist condi- tions that resulted in the southward dis- placement of a northern biota. Fossil remains of typical Hudsonian and Ca- nadian zone mammals (including various shrews, microtines, sciurids, caribou, muskoxen, moose, and the marten) have been described from Wisconsin deposits far to the south of their present distri- butions (Banfield, 1962; Barbour, 1931; Benninghoff and Hibbard, 1961; Cush- ing, 1945; Findley, 1953; Guildav, 1967; Hay, 1923, 1924; Hibbard, 1949, 1970; Hibbard and Hinds, I960; Jakway, 1958; Murray, 1957; Schultz, 1934; Schultz et al., 1951; Semken et ciL, 1964; Skinner, 1942; Stearns, 1942; Wilson, 1967). There is good evidence that during a part of each interglacial period (Aftonian, Yar- mouth, and Sangamon), subtropical cli- mates extended farther northward than now ( Hibbard, 1960 ) . Subsequent tran- sition to subhumid, mesothermal climate led to each following glacial episode. Bergmann's Rule states that within a given species of warm-blooded animal, body size increases with latitude, thus implying an adaptation that presumably serves to decrease the surface to volume ratio, thus conserving body heat in cold environments. Guilday et al. (1964), Flibbard (1963), and Parmalee (1967) presented evidence of shifts in popula- tions in some late Pleistocene faunas in response to climatic change; thus, larger northern taxa were replaced by smaller southern relatives coincident with ame- lioration of cool post-glacial climatic con- ditions. Modern representatives of these same species display a similar pattern of variation. Although details are conjectural, evi- dence of the displacement of biotic as- semblages during the Pleistocene seems impeccable. Voss (1939) estimated that the biota inhabiting four million square miles was either obliterated or displaced during this period. Amelioration of cli- mate during the interglacial stages al- 30 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY lowed migration northward; reversal of climatic trends during glaciation resulted in displacement southward. Some iso- lated relict colonies were left in favor- able or tolerable locations along the various routes of dislocation (MacGini- tie, 1959). Massive shifts of biotic assemblages occurred several times in response to changing climates and resultant glacia- tion. Although the sequence of events for each glacial episode and subsequent interglacial stage presumably was simi- lar, it is necessary to consider only the effects of the last continental glacier and ensuing post-glacial events (including additional minor oscillations) to explain the composition of the Recent biota on the Black Hills. Effects of temporary isolation imposed by Pre-Wisconsin gla- ciations would have been damped by reinvading elements through genetic swamping that would be permitted by the next advance and retreat. Much of the biota on the Hills is endemic at the subspecies and varietal levels of diver- gence. Had the current biota been effec- tively and totally isolated by Pre-Wiscon- sin episodes, endemism at the species level presumably would be more pre- dominant in the Black Hills. In addition, the warm hiatus of each interglacial stage seemingly would have affected the boreal and montane components ad- versely, causing extensive extirpation and eliminating potential centers of dispersal. Wiscomin Events. — The precise cli- matic conditions in South Dakota and Wyoming during maximal continental and Cordilleran glaciations remain open to question. Dillon (1956:168) proposed that dry, cold expanding air masses de- scended off the icecap as a result of anti- cyclonic circulation. This resulted in a mean annual rainfall of about 10 inches in the periglacial regions. However, the pollen spectrum provides evidence of a somewhat more mesic environment on the Northern Great Plains during maxi- mal glaciation. The northern ice sheet was an impediment to the southward flow of cold Arctic air onto the plains (Bryson et ah, 1970); thus, winters were warmer than at present. Absence ( or only weak development) of hot dry wester- lies, which did not begin to increase in strength and effect until the beginning of the Boreal Period ( Bryson and Wend- land, 1967), resulted in cooler, more moist summers than at present. In pos- tulating climates and life-zones coinci- dent with Wisconsin glaciation, Dillon {loc. cit.) proposed a depression (from the present) of 10-25 F in mean annual temperature for the unglaciated portion of the Northern Great Plains. His pro- posal is based on the assumption that 45^ was the mean maximum temperature under which a glacier could continue growth. Because of their higher eleva- tion, the Black Hills currently are cooler in summer, warmer in winter, and re- ceive greater annual rainfall than do the surrounding plains. Therefore, chmatic conditions suggested for the plains dur- ing Wisconsin time may be somewhat extreme for the Hills, which presumably would have been more mesic and less subject to wide seasonal fluctuations in temperature than other periglacial re- gions. Speculation has varied concerning the nature of the vegetation immediately bordering the ice sheet. Tundra (Mar- tin, 1959; Watts, 1967), grassland (Shimek, 1948; Kendeigh, 1961; Frey, 1965 ) and forest ( Clements and Chaney, 1937; Flint, 1957; Byers, 1961) have been assigned to the periphery of the ice front. Rudd ( 1951) postulated that the con- tinental plains were too cold and dry to support continuous forest during glacia- tion. The forested margins of present- day montane glaciers and the previously mentioned evidence for warmer winters and more mesic summers on the plains during maximal glaciation contraindicate Rudd's proposal. Potzger and Tharp ( 1947) and Deevy ( 1949 ) first emphasized the biogeo- graphical implications of spruce pollen occurring far south of its present distri- bution during glaciation. Additional TURNER: MAMMALS OF TliK BLACK HILLS 31 palynological studies have substantiated these earlier reports, and confirmed that a belt of fir {Abies sp.), hemlock {Tsuga sp. ), and spruce {Picea sp.) existed across the central portion of the United States during maximum glaciation. Birch {Betula sp.), alder (Alnus sp.), and tamarack ( Laiix sp. ) were present also. Sedges (Cypemceae) and sagebrush {Artemisia sp.) were the main herb con- stituents of the boreal forest (Wright, 1970). Throughout the southern states, from northern Florida to Texas, boreal plant species were present but repre- sented a more limited percentage of the total Pleistocene pollen-rain. For ex- ample, Hafsten (1964) noted that pres- ent-day grassland areas of west Texas were occupied by pine during late Wis- consin time. Recent studies (Watts and Wright, 1966; Kapp, 1970; Wright, 1970) suggested that in the Northern Great Plains, at least the Dakotas and portions of Kansas and Nebraska were occupied by a boreal spruce forest in Wisconsin time. Fossil spruce wood in glacial drift from Brookings County, South Dakota, has radiocarbon dates greater than 30,000 BP (Lemke et al, 1965:21). The current latitudinal and altitudinal limits of spruce distribution approximate the 70 F July average isotherm (Kapp, 1970). Braun (1951; 1955), Kendeigh (1961), and Thomas (1951) postulated a temperate biota inhabiting the margins of the glacier in the Great Lakes region and eastern North America, and argued for relatively little vegetational change south of the ice cap. In contrast. Gush- ing (1965) and Gurtis (1959) favored an azonal mixed coniferous-deciduous vegetational margin south of the ice front. On the bases of pollen stratig- raph\ , periglacial geomorphology, plant macrofossils, and meteorological princi- ples, other authors (Dillon, 1956; Mac- Ginitie, 1959; Martin, 1959 and else- where; Guilday et al., 1964; Bryson et al., 1970) proposed vegetational zones of transition. These transitional zones pro- gressed southward from apparently tree- less tundra to boreal woodland, decidu- ous forest in the east or pine savanna and mixed gallery forest in the west, and finally to steppe or pine savanna farther south. Pollen diagrams from artesian-spring marshes near Muscotah, Kansas, and from southern Minnesota provide no evi- dence of tundra bordering the ice sheet during the Full-glacial Period (Wright, 1970). However, pollen and plant mac- rofossils indicate a narrow zone of tundra did exist between the boreal for- est and glacial front in northeastern Minnesota at a slightly later time (Watts, 1967). The boreal flora evidently followed the glacier as it retreated northward in response to climatic change. Deteriorat- ing spruce forest and the succeeding vegetation on the Great Plains varied from east to west corresponding to changes in moisture and from north to south in relation to temperature gra- dients and shifting isotherms. The boreomontane character of the Black Hills was established during the Late-glacial times. This interval, which followed maximal advance of the Wis- consin glacial front, was a time of stag- nation, retreat, and periodic minor re- advance of the ice sheet. Presumably there was a general withdrawal of boreo- montane elements from the Northern Great Plains, accompanied by an increas- ing invasion of temperate species. Dis- junct populations of boreomontane mam- mals were left behind the retreating ice border in the Black Hills and in pockets of suitable sites across the plains. Coincident with the northeastward retreat of the continental ice sheet, the boreal spruce forest may have spread over those parts of the Northern Great Plains that previously might have been treeless. The impressive cover of paleo- sols, sand dunes and loess (Peorian and Bignell) deposited over much of the region in late Pleistocene time opposes the hypothesis of a dense periglacial forest, at least northward from western Nebraska to the glacial border (Smith, 1965; Wright, 1970). Ecogeographic dis- 32 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY tiibutional patterns of steppe-related mammals also refute the presence of a dense woodland throughout the plains at this time (Hoffmann and Jones, 1970). Schultz et at. (1951:28) suggested that the formation of loess indicated a time of "reduced vegetative cover." In north- eastern Kansas, the nature of the loess, the molluscan fauna of the loess, and the pollen diagrams from Muscotah indi- cate wooded environs. Farther west, the loess and associated molluscan fauna suggest a prairie environment interlaced with gallery forests that provided habi- tats for nongrassland species ( Leonard and Frye, 1954). Kendeigh (1961) noted that the nature of the loess of the plains was indicative of grassland abutting di- rectly against the glacial front. How- ever, meteorological evidence suggests that grassland \egetation in the Northern Great Plains during the Late-glacial in- terval may have been restricted to the more arid rain shadows of the Rocky Mountains, in western Kansas and east- ern Colorado (Bryson and Wendland, 1967). Sand dunes are not well correlated with Pleistocene events, but the Bignell Loess is radiocarbon dated at about 12,600 BP (Watts and Wright, 1966), and maximum loess deposition occurred in Illinois and Iowa from 20,000 to 14,000 BP (Wright, 1970). Smith (1965) suggested that large transverse sand dunes were formed by periglacial winds from the north, at a time when unfor- ested terrain could not interfere with dune development. Juxtaposition of the vast area of desertlikc dunes in northern Nebraska in contrast to the well docu- mented occurrence of a boreal spruce forest in other regions of the plains dur- ing Late-glacial time is compromised by Wright (1970). Retreat of the ice front, with subsequent decrease in velocity of periglacial winds, permitted the en- croachment of vegetation that stabilized the dunes. Presence of the subalpine spruce forest in the Black Hills may indi- cate that boreal vegetation then spread rapidly over (or around) the Nebraska Sand Hills, eventually reaching the Black Hills. Dillon (1956:173) mapped the hypo- thetical distribution of white spruce dur- ing Wisconsin time and suggested that boreal elements entered the Hills via a link with the Rocky Mountains. How- ever, were this the case, the species in the Hills probably would be Engelmann's spruce {Picea engelmonni) (Halliday and Brown, 1943). Thus, the white spruce in the Black Hills seemingly was displaced originally from the Cana- dian or Hudsonian life-zones to the north. Evidence of coniferous species through- out South Dakota during some part of the late Pleistocene implies that boreal components could have invaded the Hills from several directions, following cli- matic change and resultant glacial re- treat, rather than solely from the west. The present boreal elements in the Hills are disjunct from contiguous boreal for- est, which is located some 435 miles to the north (Buttrick, 1914). The cordilleran-montane biota also entered the Black Hills during the Full- glacial Period. Regional snow lines and montane biotic zones were depressed vertically about 4000-4500 feet during that time (Martin, 1959:394; Richmond, 1965:228; Webb, 1965:457). Richmond {loc. cit.) suggested a decrease of 16- 17 "F in summer temperature concurrent with the displacement. Coincident with these events, the cordilleran-montane ele- ments extended considerably downslope, eastward and southward, in\'ading the Black Hills, Laramie Mountains, and Great Plains along a wide front. Ensu- ing climatic changes during Late-glacial and Holocene times caused retreat of these components toward their previous centers, leaving relics in the Hills and other montane environs, on escarpments, and in favorable mesic sites on the plains. Montane floral elements of the Black Hills currently are disjunct from those to the west, the nearest populations residing in the Big Horn Mountains of Wyoming, about 150 miles distant. Subsequent climatic oscillations were TURNER: xMAMMALS OF THE BLACK HILLS 33 not of equal direction or intensity. The chronology of climatic events depicted in Table 2 is based on episodes defined by Reid A. Br^son and others (Biyson and Wendland, 1967; Baerris and Bry- son, 1965; Bryson et al, 1970). More recent episodes ha\'e been further clari- fied by studies of cultural history (Leh- mer, 1970) and of historical droughts (Tomanek and Hulett, 1970). Notations concerning biotic events follow Hoff- mann and Jones (1970), Watts and Wright (1966), Wells (1970a, 1970b, 1970c), and Wright (1968, 1970). Post-Wisconsin Events. — Climatic change that finally terminated the Pleis- tocene epoch brought about rapid phyto- geographic change. On the eastern por- tion of the plains, spruce forest suc- 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 other temperate deciduous elements such as elm ( Ulmus sp. ) , hop hornbeam, hazel {Conjlus sp.), and oak {Quercus sp.), with parklike openings inhabited by herbs and grasses (Watts and Bright, 1968; Wright, 1970). To the west in Kansas, Nebraska, and the Dakotas, the boreal forest gave way diiectly to prairie species (Wright, 1968, 1970). Two rec- ords of tundra-related mammals noted for South Dakota suggest an environment equivalent to present day Hudsonian and Canadian life-zones. Martin (1959:398) reported a muskox (questionably Ovi- hos) with a radiocarbon date of 9700± 600 BP, and Green and Lilligraven (1965:48) Hsted a fossil caribou from the late Pleistocene in Gregory County. Re- lict populations of boreal and montane species of plants and animals on the Black Hills also attest to these environ- mental conditions (Jones, 1964:21). Time of demise of the boreal spruce forest in the plains region has been re- corded by various authors, as follows: Muscotah, in northwestern Kansas, 15,880-11,340 BP (WMght, 1970); Rose- bud, in the Sand Hills near the Nebraska- South Dakota state hne, 12,600 BP (Watts and Wright, 1966); Rosebud in western Kansas and Nebraska, 11,400- 9100 BP (Ruhe, 1970); Pickerel Lake, in northeastern South Dakota, 10,670 BP (Watts and Bright, 1968); Pickerel Lake in Iowa, 8000-7000 BP (Ruhe, 1970). The tree-prairie transition is dated by Wright ( 1970) at 12,000 BP in the south- ern portion of the Great Plains and 9500 BP in the northern section. Later occurrence ( 5040 BP ) of wood- lands (with somewhat more pine than present today) on the plains of north- central Nebraska and South Dakota is well documented also (Kapp, 1970; Sears, 1961; Watts and Wright, 1966; Wright, 1970). Wells (1970b) recorded the mid-post-glacial flora of the Laramie Basin, southwest of the Black Hills, as being an open xerophilous woodland dominated by western red cedar and ponderosa pine with lower synusia of semidesert slnubs and grasses. The available evidence suggests a more or less azonal plant and animal community similar to that postulated by Jones (1964:23) for Nebraska during maximal glaciation in the Northern Great Plains during Late-glacial and early post- glacial times. Both prairie and forest components may have occurred in the same general area to form a savanna-Hke landscape. The relative frequency of boreal elements decreased from north to south and deciduous elements decreased from east to west. Relatively cold-toler- ant, widely distributed temperate species of grassland affinity may have been seg- regated ecologically in accordance with local conditions (Jones, loc. cit.). The supposition of an admixture of northern and southern elements in the plains re- gion also is in agreement with the view- point of Smith (1957:207). During warm, relatively moist inter- vals of post-glacial time, such as in the Sub-Atlantic or Neo-Atlantic periods, elements of the eastern deciduous forest and tall-grass prairie biotas probably extended westward far onto the plains ( eastern deciduous species via mesic gal- lery forests along river systems and val- leys, and tall-grass components on the uplands). Hardier boreal elements re- maining from the Late-glacial Period presumably could have survived these somewhat mild climatic conditions in comparatively undisturbed, disjunct pop- ulations. Jones (1964) interpreted the relatively large amount of fossil pine pollen described from the Nebraska Sand Hills by Sears (1961) to indicate that boreal species survived early post-glacial time only to be excluded by ensuing mid- post-glacial periods of maximum warmth and dryness. The eastern deciduous bi- ota may have mingled with residual cor- dilleran elements (from Pleistocene or Sub-Boreal times when the biota spread eastward and southward down the slopes of the Cordillera) at points throughout the Great Plains (Webb, 1965). Conversely, arid post-glacial periods TURNER: MAMMALS OF THE BLACK HILLS 35 were a time of extirpation o£ the boreal and eordilleran-montane elements that survived from late Pleistocene time on the Northern Plains; only the most tol- erant species survived in small, isolated populations. Much of the unique biota of the Black Hills may have been re- stricted during these periods. Outlying representatives probably could not sur- vive the effects of the hot, dry climate; the milder, more stable climate in the Black Hills presumably afforded a suit- able refugium for these elements. Those species of eastern and southern affinities that had spread into the Northern Great Plains area during the humid climatic episodes either found suitable refugia during the time of exapotranspirative stress, or were subsequently excluded. During the mid-post-glacial time of maximal aridity, xeric plant species sup- planted mesophytic vegetation; these species fragmented formerly continuous areas of deciduous vegetation and per- mitted encroachment of arid-tolerant trees and nonarboreal plants (Smith, 1965). Gleason (1923) invoked an east- ward extension of the prairie to explain the relict pattern of prairie plant distri- bution (prairie peninsula) in the Mid- west, and postulated the extension prior to the invasion of deciduous species in post-glacial time. Transeau (1935) in- dicated that the extension was subse- quent to the initial invasion of the decid- uous forest in mid-post-glacial time. Available evidence implies that maximal eastward penetration of the grassland occurred in the Atlantic Period. How- ever, as with the north-south glacial dis- placement of the biota, the west to east dispersal of prairie species may have been repeated several times. Thus, during milder intervals of post- glacial time, species having eastern and possibly southern affinities spread onto the Northern Great Plains and Black Hills, and mixed with residual boreal and eordilleran-montane constituents. During subsequent arid climatic epi- sodes, these immigrants either inhabited suitable refugia or were eliminated when drought environment promoted an east- ward expansion of the steppes. In time, amelioration of dry conditions resulted in retreat of grasslands to their present limits. This sequential series of events can be traced in pollen diagrams of Pick- erel Lake and Muscotah (Watts and Bright, 1968; Wright, 1970). Wells (1970b) described the present flora of the Laramie Basin as desertic in physiog- nomy with strong resemblance to winter- cold deserts of the Great Basin and Colo- rado Plateau. Change from a xerophi- lous woodland to a semidesert shrubland in post-glacial time denotes a secular trend toward an increasingly arid climate east of the Laramie Front Range. Re- cession of the boreal forest barrier that existed between the Great Basin and Wyoming Basin during the Wisconsin ( Hoffmann and Jones, 1970 ) opened ac- cess for cold-desert species to the North- ern Great Plains. Shifts in upper-air anticyclonic circulation northeastward from the Great Basin in the Sub- Atlantic and resultant influx of tropical air also may have influenced the biota of the region. For example, evidence of exten- sive introgression between the bur oak in the Black Hills and Gambel's oak ( Qiier- cus gambelii) to the southwest, in the central Rocky Mountains, presupposes a massive northeastward migration of the cold-intolerant Gambel's oak during a warm moist period ( Wells, 1970b ) . Such conditions were not current during east- ward and southward displacements of eordilleran-montane elements. The sequence of episodes in the his- tory of native cultures of the Missouri River Valley in the Dakotas (Lehmer, 1970) and the Mill Creek culture of Iowa (Bryson et al., 1970) implies a close cor- relation between climatic and cultural changes in near-modern time. Modera- tion, culminating in the present climatic conditions, has been interrupted by a gradual warming trend on the North American continent in the last 100 years (Dorf, 1959, and others). Periodic, ex- tended droughts have resulted in changes of the grasslands and associated fauna 36 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY (Tomanek and Hulett, 1970). Wooster (1935, 1939) and Gier (1967) noted that some mammals, such as the prairie vole (Microtiis ochrogaster) , almost disap- pear in some areas under arid regimes, whereas others, such as the deer mouse (Peromijscus maniculatus) , seem rela- tively unaffected by extended drought. The Great Plains of Recent time is an extensive grassland on deep, trans- ported soils, corresponding to flat or gently rolling topography. The gram- inoid flora is defined by a predominance of a few widespread species and a pau- city of endemics, virtually none of which are grasses (Wells, 1970c). The associ- ated insect fauna (e.g., range grasshop- pers and lataline grass-feeding leafhop- pers), has similar distributional charac- teristics; however, the fauna has a much greater species diversity in certain serai stages of surrounding forested biomes than in comparable grassland communi- ties (Ross, 1970). Generally, trees are re- stricted to riparian habitats as gallery forests along water courses; in addition, nonriparian woodlands occur on escarp- ments and other topographic breaks throughout the grassland province of central North America. Clements and Chancy (1937) indi- cated that the prairie species probably developed after Miocene time in the central parts of the continental United States due to drier conditions created by mountain building to the west. How- ever, Wells (1965, 1970a, 1970b, 1970c) and Ross ( 1970 ) presented several argu- ments for relatively recent tenure of the central grasslands. Accumulated evi- dence indicates that these extensi\'e tree- less steppes probably have not had a con- tinuous existence since the mid-Tertiary. The abundant, nonarboreal pollen re- ported by Kapp (1970) from interglacial sediments of the Sangamon interval may be from restricted refugia; the occur- rence of the pollen does not necessarily imply the presence of widely distributed steppe vegetation. During maximal Wis- consin glaciation, grassland species sur- vived in suitable refugia southwest and southeast of the glacial front or possibly as mosaic distributions in a prairie-forest savanna. When one realizes that most grassland species have a major part of their diverse ranges as synusial compo- nents of serai stages of forested en\'irons, the uniqueness of the prairie biome be- gins to fade; consequently, the grasslands should be regarded as consisting of a derivative biota. Climate alone does not account for a grassland climax vegetation. In areas where grasslands were planted in Ne- braska with upland tree species native to scarp woodlands, rather extensive for- est became established (Wells, 1965). Furthermore, Potter and Green (1964: 22) found evidence of ponderosa pine, as seedlings and young saplings, invad- ing the grasslands in North Dakota. A similar invasion can be observed on the prairie of Wind Cave National Park, where grazing by bison checks the re- production of ponderosa pine. Thus, in at least certain locations, the plains are quite capable of supporting nonriparian woodlands (at least of xerophytic coni- fers). Physiography may be as impor- tant a factor as climate in affecting the distribution of extensive, treeless grass- lands. Stewart ( 1951 ) noted that theoretical climatic climax vegetation in savanna and grassland areas probably postdates the arrival of man. The Tule Spring, Nevada site, dated at +23,000 BP, clearly indicates that prehistoric man was present during maximum glaciation in the Wisconsin (Martin, 195S). The fre- quency of fires, which generally is as- sumed to have increased with the arri\ al of man, is well documented in early his- torical literature. Scarps and abrupt topographic breaks may have ser\ed as refugia from grass fires for the nonri- parian \\'oodlands of the plains region (Wells, 1965, 1970a, 1970b, 1970c). Forest fires may ha\'e terminated broad expanses of boreal forest that were already suffering evapotranspira- tive stress from the effects of warm post- glacial climate. These newly opened TURNER: MAMMALS OF THE BLACK HILLS 37 expanses subsequently were invaded by a biota ha\'ing strong ecological affinities with nearby forested biomes. Interaction of topography, climate, and subsequent prairie fires apparent!)' has resulted in the present ph\ siognonn' of the Northern Great Plains. See the several papers of Wells cited above for additional argu- ments concerning these points. The introduction and development of modern ci\ilization and agriculture in the plains region no doubt ha\'e resulted in extirpation of several species, endan- gered others, and caused widespread, drastic modification of habitats that frag- mented the ranges of yet other species (Smith, 1965). In addition, man has in- troduced many exotics (e.g., Kentucky bluegrass) and thereby possibly caused the exclusion of native species or shifts in their respective ranges. The Black Hills flora is diverse, with inany species occupying a small geo- graphical area. For example, Froiland (1962) identified 20 kinds of willows (Salix sp.) of diverse geographic origin in the Hills region. This diversity is due partly to the varied topography and hab- itats available, and partly to the past movement of great floral assemblages during late Tertiary and early Pleistocene times (Wetmore, 1968). However, the Black Hills are not unique in having dis- tributional limits of so many northern, southern, eastern and western species. The probable geographic origin of the vegetative components of the Black Hills are listed in Tables 3 and 4. The presence of many widespread species (22-34%) can be attributed to the fact that plants with wide ranges of tolerance overlap distributionally within the Hills region. Most northern or boreal (6%) and western or montane (25-30%) spe- cies were displaced southward or east- ward by advance of continental and Cordilleran ice sheets, respectively. These species became stranded on the Hills by migrating up the slopes after glacial retreat; although many species presumably were extirpated in the At- lantic Period, some remain currently. Table 3. — Distril:)utional patterns of the flora of the Black Hills, as noted liy Hayvvard ( 1928) andMcIntoch (1931). Biographic origin Percent affinity Western 25-30 Widespread 22-34 Great Plains 17-26 Eastern 9 Northern 6 Southern 4—5 Table 4. — Distributional patterns of lichens of the Black Hills as noted by Wetmore (1968). Biogeographic origin Percent affinity Arctic-Boreal 44.4 Pan Boreal 24.9 Pan Temperate 5.4 Western Temperate 4.9 Arid Southwestern 4.4 Pan North America -^ 3.9 Southern Rockies- Alleghenian-Great Lakes 3.9 Eastern Temperate 3.4 Eastern Boreal 2.9 Grassland 1.5 Western Boreal 0.5 Examples of boreal species are paper birch, twin-flower, Venus' slipper, bunch- berry, squashberry, and white spruce. Montane species include Oregon grape, Indian paint-brush, mariposa lily, skunk- bush sumac, buffaloberry, white spruce and ponderosa pine. Fernald (1935) suggested that a few western species — for example, gland stem and huckleberry — may represent pre-Pleistocene floral elements. Great Plains components (17-26%) are to be expected because of the loca- tion of the Black Hills, which are entirely surrounded by plains. These plants are found in the foothills, in the drier mead- ows, and open parklands. Examples are blue grama, Indian grass, porcupine grass, and western sandcherry. Eastern species (9%) presumably reached the Hills during the Sub-Atlantic or Neo- Atlantic intervals via gallery forests along water courses and uplands. These may be Ozarkian and Alleghenian elements of the eastern deciduous forest (Braun, 38 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 1947). Examples are hop hornbeam, American elm, boxelder, Solomon's seal, green ash, and bur oak. The southern components (4-5%) probably arrived at about the same time as did the eastern elements. In part these are derived from the old Madro-Tertiary flora that developed in the southwestern United States in Miocene, Oligocene, and Pliocene times (Wetmore, 1968). The driest areas of the foothills now are occupied by the following species: soap- weed, brittle pricklypear cactus, skunk- weed, scarlet gaura, silver sagebrush, and wormwood. The examples used in this resume were taken from Wetmore (1968). The distiibutions of lichens in the Black Hills are vastly more complex than the respective distributions of vascular plants because of the diversity of sub- strates, exposed geological materials, ele- vations and relief, climatic range, and vegetational types. The probable geo- graphic origins of the lichens in the Black Hills are listed in Table 4. For a thor- ough analysis of the complexities in- volved in distributional patterns of lich- ens, see Wetmore ( 1968) . INFLUENCE OF MAN ON THE ENVIRONMENT The Black Hills region is one of the most densely populated areas of the Great Plains. In 1960, the Hills area con- tained an average of 6.2 persons per square mile. Of the native population, 57 percent were classified as urban, 30 percent as nonfarm rural, and 13 percent as farm rural. The presence of about 6000 armed forces personnel and from four to five million annual tourists inten- sifies utilization of Black Hills environ- ments. Changes wrought by man have been detrimental to some mammalian species and advantageous to others. Large herbivores and carnivores, such as the bison and wolf ( Canis lupus ) , have been directly extirpated by man; populations of other species such as the mountain lion {Felis concolor), black bear {Ursus americanus), and prairie dog, have been reduced drastically. When European man first moved into the Black Hills, big-game mammals, fur-bearing mam- mals, and smaller game mammals were abundant, but subsequently they have become the object of exploitation. As agricultural endeavors in the region in- creased, carnivores and rodents were poisoned indiscriminately. Fortunately, recent programs of wildlife management and a more intelligent application of rodent and predator control have less- ened the devastation by man. Estab- lishment of Wind Cave National Park, Custer State Park, and several other areas has served to protect and pre- serve remnants of the native fauna. Man has reintroduced several species that for- merly were native to the Black Hills, such as the elk (Cervus canadensis), pronghorn, bison, and mountain sheep. In addition, the mountain goat, fox squir- rel (Schirus niger), house mouse {Mus muscuhis) , and Norway rat (Rattus nor- vegicus), which are not native to the Hills, were introduced by man. Activi- ties of man that have indirectly affected the abundance and distribution of mam- malian species in the Black Hills are summarized below. For detailed infor- mation concerning these actixities, the reader is directed to the "Black Hills Area Resource Study," published jointly by the Departments of Agriculture and the Interior in February 1967. Mining. — The gold rush of 1874-79 gave impetus to the mining industry in the Black Hills region. In the early 1900's, mining of silver, tin, coal, and crude oil flourished. The two World Wars created a greater demand for vari- ous minerals and gave added impetus to the tungsten, sheet mica, feldspar, ben- tonite, beryl, and lithium industries in the Hills. The most recent mining boom began in 1951, with the discovery of uranium ores in Jurassic-Cretaceous sandstones. Production of nonmetallic commodities, such as sand, gravel, crushed rock, cement, and clay, also are current industries of the region. Mining TURNER: MAMMALS OF THE BLACK HILLS 39 in\'aiiably alters the natural environment. For example, in processing bentonite, the overburden is stripped off and the bentonite is turned oxer and stored in place for long periods of time. Excava- tion of mining complexes and rock quar- ries also has altered the topography. Processing of gold in the Lead-Dead- wood area has resulted in pollution of man)' nearby streams. However, not all alterations ha\'e been detrimental to the regional fauna; for example, mines fre- quently are utilized by woodrats and various species of bats. Timherinp,. — Cutting of timber com- menced simultaneously with the Black Hills gold rush, and at least two portable, steampowered sawmills existed in the Hills by 1877. Intensive timbering caused a rapid depreciation of the quality in native stands adjacent to mining opera- tions by 1897, when President Cleveland created the Black Hills National Forest Reserve. Sale of federal timber was authorized by the Timber Reserve Bill, enacted later in the same year. The timber industry developed rapidly, sup- plying vast quantities of wood products to the mines, railroads, and other indus- tries of the region. The pulp and paper, and the post and pole industries have flourished recently. Forest production accounts for about 14.7 percent of the principal land use in the Black Hills. The total of commercial timber currently available in the Hills is estimated at five billion board feet; production in 1964 was in excess of 21.8 million cubic feet, contributing 6.5 million dollars to the regional economy. Exploitation of for- ests by lumbering is evidenced by vast areas of scrubby second-growth timber and numerous open areas where burning has resulted from lumbering operations. Removal of dead trees in clearing opera- tions has eliminated potential den sites for flying squirrels and red squirrels. However, populations of some mammals (e.g., deer and chipmunks) have in- creased in areas of stumps, logs and trim- mings. Agriculture. — Homesteaders first moved into the Black Hills about 1900, but the drought of 1910-11 shifted major emphasis from general farming to the raising of lixcstock. Ranching became an important activity in the unforested uplands, foothills, and the Red Valley. Grazing now accounts for 75.1 percent of the principal land use in the Black Hills area; usually beef cattle and sheep are marketed as feeder stock. About 103,000 acres were under irri- gation in the Hills region in 1960. Crop- land comprises about 8.6 percent of the principal land use, with corn, sorghum, winter wheat, spring wheat, barley, al- falfa, and hay being the major dryland crops. The average farm in the Hills area in 1960 was 2550 acres in extent, and sales of agiicultural products amounted to more than 50 million dol- lars. The en\ironment has been modi- fied conspicuously due to agricultural practices. The original prairie grasslands of the Red Valley have been altered by gi'azing and the upland grasslands (e.g., the Reynolds and Gillette prairies) now support hayfields, market gardens, and pastures. Wind Cave National Park is one of the few large areas wherein native prairie grassland still exists nearly in its original state. Fires. — The earliest records of fire in the Black Hills were about 1730 to 1740, when the entire area appears to have burned by a series of fires; in the period between 1790 and 1800, most of the Hills were burned again (Graves, 1899). Lesser fires of considerable extent oc- curred in 1842, 1852, and 1875. From 1880 through 1966, at least 70 fires in the Black Hills burned more than 175,000 acres of forest and grassland. Of these, 44 percent were known to have been caused by man and 18 percent by light- ning, whereas 38 percent were of un- known cause (records of U. S. Forest Service, Black Hills National Forest). Among the more notable fires were those of Iron Creek in 1899 (38,000 acres), Rochford in 1931 (21,640 acres), McVey in 1939 (21,875 acres), and Wildcat in 40 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY 1960 (10,336 acres). Improved precau- tionary measures and modern fire-fight- ing techniques have prevented major fires in recent years. Two smaller fires should be mentioned. Although the Deadwood fire burned only 4501 acres, it forced evacuation of that historic town in 1959. The Headquarters fire in Wind Cave National Park in 1964 (4000 acres) greatly endangered the natixe wildlife in the Park. A noticeable result of forest fires is the opening of extensive new grassland, and the establishment of as- pen and birch woodlands in burned areas. Reforestation programs and nat- ural secondary regeneration have par- tially revegetated the scarred hillsides that resulted from past fires. Recreation and Tourism. — Although immeasurable in extent, perhaps the most detrimental influence on the environ- ment of the Black Hills in the past few decades has been the influx of four to five million tourists annually. These visi- tors have intruded into canyons, onto the forested slopes, and upon much of the remainder of the relatively undis- turbed en\'ironment in countless \\'ays. Recreation and tourism are the foremost producers of revenue in the Black Hills; more than 120 million dollars were con- tributed to the regional economy by this industry in 1966. People seeking recrea- tion are attracted to the Black Hills by points of historic and geologic interest, by opportunities to hunt and fish, and by such scenic features as Custer State Park, Wind Cave National Park, Jewel Ca\'e National Monument, Devils Tower National Monument, and Mount Rush- more National Memorial. Demands for recreation surely will continue to in- crease in the future, as will exploitation of the once pristine environment. ACCOUNTS OF SPECIES ORDER INSECTIVORA— Insectivores Family SORICIDAE— Shrews In the Black Hills, the order Insecti- vora is represented by one species of the genus Sorex. An additional species of this genus may occur in the Hills, and another has been incorrectly reported from the area. Sorex cinereus haydeni Baird Masked Shrew Sorex haydeni Baird, 1858, Mammals, in Re- ports of explorations and surveys . . . from the Mississippi Ri\er to the Pacific Ocean . . ., 8 (1):29, 14 July (type locality re- stricted to Fort Union, just west of conflu- ence of Missouri and Yellowstone rivers, Williams Co., North Dakota, by Merriam, N. Amer. Farma, 10:60, 31 December 1895). Sorex cinereus haijdcni — Jackson, 1925, Jour. Mamm., 6:56, 9 February. Vernon Bailey (1888:436) first re- ported the masked shrew from the Black Plills under the name Sorex personatus. Known altitudinal range in the Hills ex- tends from 4500 to 6500 feet, but this shrew is most widely distributed in the Central Basin and on the Limestone Plateau above 5500 feet. It is common locally in riparian associations and other moist habitats throughout the Northern Great Plains. Except for three individ- uals ol:)tained in October and November, all specimens examined were taken in warm months. In the Hills, this species is most nu- merous in montane habitats; it is locally abundant in marshy areas, mossy bogs, and other riparian associations of grasses, sedges, rushes, aspen, birch, and \\ illow. Many specimens were trapped near logs, large rocks, or low shrubs located in dense grass along creeks such as Beaver, Boxelder, Castle, Cold, Grizzly, Iron, Rapid, Rattlesnake, Spring, and Willow, or in the xicinity of beaver dams. Spring- fed bogs and other moist areas in aspen and spruce woodlands also pro\'ide suit- able habitat for these shrews. The availability of ground water un- doubtedly is one of the most important factors limiting the distribution of S. c. haijdcni. A few indi\iduals ha\e been taken in unusual habitats; for example, TURNER: MAMMALS OF THE BLACK HILLS 41 a female was trapped on a rock ledge near the top of a 30-foot cliff, and four others were obtained on dry, rocky, pine- clad hillsides approximately 200 feet from the nearest source of water. Also, two shrews taken by Bailey (1888:436) were captured "at holes in the rocks on top of one of the highest peaks." Popu- lation densities frequently are low in these marginal areas. Associated with the masked shrew in moist habitats are Zapus hudsonhis, Microttis longicoudns, and M. penn.syhanicus. Feromijscus maniculatus, Chthrionoimjs gapperi, and Eufaniia.s luinimtis have been taken with these insecti\ores in drier environments. A member of a University of Kansas field party, J. R. Choate, buried 20 cans in tall grass along Boxelder Creek, 20 mi. W Nemo between 29 June and 5 July 1967. The cans were buried 10 paces apart with the openings at ground level. A snap-trap was placed within five feet of each can. These pitfalls (three shrews captured) seemed to be a more eflBcient method of obtaining S. c. haydeni than was snap-trapping (one shrew cap- tured); however, these data are insuffi- cient and a more thorough application of this technique in the Hills region is desiralile. Trapped insects may have attracted the shrews to the pitfalls. Reproductive data are a\ailable for S. c. haydeni in the Black Hills from mid- June through mid-July. Of 33 females captured during this period, 15 showed no sign of reproductive activity; the same was true for a female taken in Spearfish Canyon on 7 August. Mammae of a fe- male obtained on 18 June were enlarged, whereas another captured on 24 June had an enlarged uterus, possibly indicat- ing recent implantation. Twelve females taken in the period 25 June to 7 July carried 5.2 (2-7) embiyos that were 5.9 (2-11) in crov-'n-rump length. Four lac- tating females were noted in the period 27 June to 13 July. The testes of 14 males captured early in the summer were 4.5 (2-6) in length, whereas those of three taken earh' in August were 3. Pruitt ( 1954 : 36 ) reported that reproduction oc- curs only in fully matured or old individ- uals in S. c. cinereus in Michigan. Three young females of age class two of Pruitt ( loc. cit. ) were obtained in summer and were gravid; these were taken on 25 June (seven embryos), 29 June (five em- bryos) and 2 July ( five embryos ) . Pruitt also indicated that the testes of reproduc- tively acti\'e males exceeded two milli- meters in length; testicular length of an immature male obtained on 3 July was 5. While uncommon, such precocious repro- ductive activity has been reported in other species of the genus Sorex. Young of the year were evident from late June through mid-August. Due to the uniformity in season of collection (mid-June to early August), most of the specimens examined were in summer pelage. However, a male ob- tained by Vernon Bailey near Sundance on 13 August 1913 evidenced molt, with fresh winter pelage covering about two- thirds of both dorsum and venter; three others, taken by Merritt Cary in the Bear Lodge Mountains on 22 June 1912, had just completed molt. In considering geographic variation, specimens of S. c. hoydeni from the Black Hills were compared with individ- uals from south-central Wyoming (Al- bany and Carbon counties). North Da- kota (Bowman and Walsh counties), eastern South Dakota (Kingsbury, Union, Marshall and Day counties), and northern Nebraska (Sioux, Sheridan, Cherry, and Keya Paha counties). Cra- nial measurements were taken in the manner described bv Jackson (1928:12) and Findley (1955:5). Cranial breadths of 29 females averaged somewhat greater (7.8±0.21) than those of 22 males (7.6 ±0.22). Otherwise, significant secondary sexual differences were not apparent nor was significant variation within the Hills population detectable. Reflectance readings show that masked shrews vary geographically in tone of color. Individuals from North Dakota and north-central Wyoming a\'er- aged paler than those from the Black Hills, wliereas specimens from south- 42 MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY central Wyoming and Nebraska aver- aged darker than Hills specimens. Shrews from eastern South Dakota ap- proached those from the Hills in colora- tion ( Table 5 ) . No significant geograph- ical difl^erences in hue were discernible. Both external and cranial dimensions evince considerable geographic variation. Masked shrews from localities west of the Black Hills have greater average body, tail, maxillary tooth-row, and pal- ate lengths, as noted by Long (1965: 519). However, these individuals are indistinguishable from eastern represent- atives on the basis of condylobasal length and interorbital breadth. In addition, the average rostral length is greater and the breadth across the maxillary proc- esses is significantly less than noted in specimens from sites east of the Black Hills. I am in agreement with Long (loc. cit.), who assigned masked shrews from northeastern Wyoming to S. c. hay- deni and those from elsewhere in that state to S. c. cineretis. In most dimen- sions, specimens from the Black Hills are intermediate between S. c. cinereus to the west in Wyoming and S. c. haydeni to the east in the Dakotas and Nebraska, but fall within the range of \'ariation of the eastern specimens and hence are assigned to S. c. haydeni. Additional measurements (not listed in table 5) of 67 adult shrews from the Black Hills are: percent red reflectance, 54.3±0.52; percent green reflectance, 23.7 ±0.19; percent blue reflectance, 21. 3 ±0.32; hind foot length, 11.2±0.64; ear length, 6.4± 1.25; weight, 3.9±0.94; condylobasal length, 15.5±0.28; cranial breadth, 7.7 ±0.23; interorbital constriction, 2.8± 0.11; skull depth, 4.7±0.33. Specimens examined (136).— SOUTH DA- KOTA: Lawrence Counti/: Deadwood, 3 (USNM); Dumont, 6100 ft, 3 (USNM); 2 mi S Tinton, 6100 ft, 6; Big SpearBsh Canyon, 9 iTii S, 3 mi W Spearfish, 5000 ft, 2; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, .5800 ft, 2; 3 mi W Nemo, 4800 ft, 2; 2 mi W Nemo, 4700 ft, 20; Nemo, 4700 ft, 1. Pc7inington County: 3 mi W Rapid City, 1; 20 mi N Elk 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. Number and sex of specimens averaged i-i o c u H o o c f«^ ^ be t? bO "rt C o c H^ p; i> ■£ J V bcT:: ^ (D rt O 4(2,^,29) Mean 28.0 86.0 SD ±4.24 ±5.66 7(55,29) Mean 20.4 SD ±5.08 13(6<5,79) J Mean 16.8 91.8 SD ±2.55 ±3.77 67(305,379) S. Mean 19.6 SD ±2.95 20 (95,119 ) Mean 22.9 SD ±3.12 73(445,299) Mean 19.0 SD ±2.75 S. c. hat/deni, North Dakota 35.6 5.1 4.3 6.0 ±0.00 ±0.07 ±0.14 ±0.14 S. c. hai/deni, eastern South Dakota 87.3 33.0 5.3 4.2 6.1 ±1.86 ±2.45 ±0.15 ±0.18 ±0.07 S. c. hatfdeni, northern Nebraska 33.5 5.3 4.2 6.1 ±2.29 ±0.12 ±0.11 ±0.13 c. hai/deni, Black Hills, South Dakota and Wyoming 92.3 37.2 5.3 4.2 6.2 ±5.14 ±3.30 ±0.14 ±0.11 ±0.16 S. c. cineretis, north-central Wyoming 93.3 37.1 5.4 4.1 6.3 ±5.54 ±2.16 ±0.12 ±0.10 ±0.16 S. c. cine reus, south-central Wyoming 96.3 38.8 5.4 4.1 6.2 ±5.50 ±2.50 ±0.15 ±0.13 ±0.16 5.4 :0.21 5.5 :0.09 5.6 :0.13 5.6 :0.14 5.7 :0.14 5.6 :0.13 TURNER: MAMMALS OF THE BLACK HILLS 43 City, 6200 ft, 1; Ditch Creek, 14 mi W Hill City, 6400 ft, 1; Palmer Culch, 3 mi SE Hill Citv, 53-5400 ft, 16 (U\LMZ); 3 mi SSE Hill City, 1 (UMMZ); Willow Creek, 4 mi SE Hill Citv, 53-5400 ft, 12 (UMMZ); Castle Creek, 6500 ft, 3 ( UNLMZ); Sprins? Creek, 2 mi W OreviJle, 5500 ft, 3 (UMMZ); Rapid Creek, 1.5 mi W Rochford, 1 (UMMZ); Bea\er Creek, 4 mi N, 10.5 mi \V Deerfield, 6400 ft, 6; 3 mi N, 7 mi W Deerfield, 6400 ft, 1 ; 3 mi S, 1 mi W Rockerville, 1; 16 mi NW Custer, 1 (UMMZ). Cttsicr Countii: 0.5 mi E Sylvan Lake, 6250 ft, 2 (UMMZ); 5.75 mi N, 5.75 mi E Custer, 5220 ft, 19; 16 mi W Custer, 2 (USNM); Custer, 3 (2 USNM, 1 AMNH); Lightning Creek, 8 mi SW Custer, 5100 ft, 1 (UMMZ). WTONHNG: Crook County: Beaver Creek, Bear Lodge Mountains, 6.5 mi SSE Alva, 1 (UMMZ); Warren Peak, Bear Lodge Moun- tains, 6000 ft, 4 (USNM); Sundance, 3 (USNM); Rattlesnake Creek, 6000 ft, 1 (USNM); 3 mi NW Sundance, 5900 ft, 10. Weston County: 1.5 mi E Buckhorn, 6150 ft, 2. Additional record?.— SOUTH DAKOTA: Lawrence County: Spearfish (USBS files); Rochford (USBS "files). Custer County: Bull Springs, 2 mi N, 9 mi W Custer (Bole and Moulthrop, 1942:95). County unspecified: Black Hills (Bailey, 1888:436)." ORDER CHIROPTERA— Bats Family VESPERTILIOxMDAE— VeSPERTILIOxNIDS Ten species of bats, representing five genera, are known to occur in the Black Hills of South Dakota and Wyoming. One additional species of another genus {Euderma) is unreported as yet, but may inhabit the area; another species has been erroneously reported from the Hills and the status of still another remains uncertain. Chiropterans from the Black Hills represent but one family, Vesper- tilionidae, in the suborder Microchirop- tera. Three species are known to migrate southward during the colder months, whereas six are resident year-round in the area; the seasonal distribution of one species (Myotis keenii) is in question. The Black Hills contain a large num- ber of potentially favorable retreats for many kinds of bats. The mountainous ten-ain is comprised of uplifted segments of granite, and exposed cliffs and rock ledges of limestone and sandstone. Huge caverns and lesser caves have formed naturally in the soluble limestone layers. Jewel Cave, Wind Cave, Davenport Cave, Ice Cave, Bear Trap Cave, Igloo Cave, and the numerous unnamed and/ or commercial caves of the region serve as hibernacula in the colder months. Mining in the Hills began with the gold rusli of tlie mid-l', 6200 ft, 7; Porcupine Draw, 14 mi W' Hill City, 6400 ft, 1; Ice Cave, 8 mi N, 15 mi W Custer, 6400 ft, 9; T. 2 S, R. 6 E, E 'A sec. 19, 1 (MS\VB); 10.5 mi E Wyoming border, 5.8 mi N Custer County line, 8 (MSWB). Custer County: Elk Mountain, 1 (USNM); Jewel Cave, 2.5 mi S, 12 mi W Custer, ,5280 ft, 10 (2 SDMT; 8 UK); Custer State Park, 2 (UMMZ). WYOMING: Weston County: 0.5 mi E Buckhorn, 6100 ft, 3. Myotis leibii ciliolabrum (Merriam) Small-footed Myotis Vespertilio ciliolabrum Merriam, 1886, Proc. Biol. Soc. Washington, 4:2, 17 December ( type locality, a bluff on Hackberry Creek, about one mile from Castle Rock, near Banner, Trego Co., Kansas). Myotis leibii ciliolabrum — Glass and Baker, 1965, Bull. Zool. Nomenclature, 22:204-205, August. The small-footed myotis is known from all five counties that comprise the Soutli Dakotan portion of the Black Hills, but it has not been reported previously from northeastern Wyoming. This bat, which has been taken in the Hills year- round, is known to hibernate in Jewel Cave, Davenport Cave, and in an un- named limestone cave in Dark Canyon, Pennington County. Robert A. Martin (pers. com.) recently has located the 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 Tone of color Percent red reflectance Percent green reflectance Percent blue reflectance 19 (14^,59) Mean SD Black Hills, South Dakota and Wyoming 16.4 54.2 24.3 21.5 ±1.77 ±0.22 ±0.19 ±0.17 4(25,2$) Mean SD Bon 19.4 ±1.65 Honune Countv, South Dakota 53.0 ' 25.7 21.3 ±0.20 ±0.17 ±0.13 6(35,35) Mean SD 17.7 ±4.73 Sarpy County, Nebraska 56.3 24.6 ±0.40 ±0.21 19.1 ±0.22 9(35,69) Mean SD Boom 15.2 ±1.77 ', Hardin, Keokuk counties, 55.5 25.2 ±0.25 ±0.13 Iowa 19.3 ±0.25 3(25,1?) Mean SD 16.7 ±2.31 Marshall County, Kansas 55.8 23.2 ±0.46 ±0.16 21.0 ±0.42 species in additional hibernacula such as the Cleopatra Mine, S and G Cave, French Creek Cave, and Igloo Cave. Al- titude does not seem to be limiting inas- much as Myotis Jeilni occurs throughout the Hills up to at least 6500 feet. It was reported first from the region by Miller and Allen (1928:169). On 2 July 1965, a horizontal shaft in the deserted Cambria Coal Mine com- plex north of Newcastle, Wyoming, was netted and yielded five M. leihii and one M. thysanodes. Two more leihii were shot over a nearby stock pond on the fol- lowing evening and another was netted in the mine shaft. On 19 November 1967, a single male was found hibernating in a cave in Dark Canyon (see account of L. noctiva