UMl
HARVARD UNIVERSITY
LIBRARY
OF THE
Museum of Comparative Zoology
JM
13
?^ 0 Not Remove
^ rnet Archive
Do Not Remove
Internet Archive Boston Scanning Center
Shipment ID _Harvard MCZ 2/13.
Item #s
This bound volume contains more than one item.
Divide here for
l5o>l
See loader or manager w/ any questions
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_^_
""<f f S j-^y"' CANYON
DEE»F,HD BmO HIILS - '""
'' " -'DEERtlELD RES SHERIDAN
DITCH CREEK CG i m,\/et burn iake ,
1 M.VEYEIURN «^^ ^^jjjUHII^
eiLLEIIf HILL CITY^^ ^.•-ROCKERVllLE
rmiNIt p,^„j, 5„^(,„ ^ KEYSTONE
HORSEIHIEF i_;-i \0
HARNEY PEAK < \mt RUSHM„„.
CUSTER CO. "f 5^ Jl°^."' "^~^
I r "■ '
ANGOSTURA RESERVOIR
EDGEMONT
44^
104°
Fig. 2. General geography of the Black Hills, showing major collecting sites mentioned in te.\t (modified after Pettingill and Whitney, 1965:frontis).
16
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
104°
104"
■15J Tertiary igneous intrusions I I Tertiary sediments
Paleozoic sediments
■■'■w'.l Precambrlan granitic rocks
. Cretaceous sediments Triassic and Jurassic sediments
Precambrlan metamorphic rocks
Fig. 3. Generalized geologic map of the Black Hills area (modified after Black Hills Area Resource Study, 1967:18).
TURNER: MAMMALS OF THE BLACK HILLS
17
shale (Opcche Formation). Along the base of the foothills lies a conspicuous, and remarkabK' continuous valley in the form of a "racetrack," \\'hich encircles the Hills (Fig. 7). This narrow trough rep- resents the lowest elevation in the Hills (3200 to 3500 feet above sea level). It cuts into soft red shales and sandstone of the Spearfish Formation, which was formed as a deposition of sediments early in the Triassic. The Red Valley, so named because of reddish soils of the Spearfish-Ne\'ille Soils Subassociation, varies in width from a quarter of a mile (in the east) to more than six miles (in the northwest); the xalley floor is dis- sected by many drainage divides that are separated by broad alluvial flats.
The Limestone Plateau, Central Ba- sin, and peaks of the Bear Lodge Moun- tains form the so-called "boreal-cap" of the Black Hills; the latter two areas comprise the Mountainland Soil Asso- ciation, whereas the former consists of the Limestone Plateau Soil Association. The boreal-cap is characterized by a pre- dominantly coniferous forest fauna of boreomontane and cordilleran origin. Mammals such as Nuttall's cottontail (Sylvilagus nuttallii), the least chipmunk {Eutamias minimus), yellow-bellied marmot (Marmota flaviventris), red squirrel (Taniiasciunis hudsonicus), northern flying squirrel (Glaticomys sa- ])rinus), northern pocket gopher {Tho- momys talpoides), red-backed vole (CletJirionomys gapperi), and long- tailed \'ole ( Microtus Jongicaiidtis), reach their highest population densities in these areas. In the Central Basin, isolated grasslands of the Slate Prairie Soil Asso- ciation ( Fig. 11 ) are inhabited mainly by the white-tailed jackrabbit (Lepus toicn- semlii), thirteen-lined ground squirrel {Spennophihis tridecemlineatus) , deer mouse {Peromysctis maniculatus) , and meadow vole {Microtus pennsylvani- cus). The crystalline core near Harney Peak provides the restricted range of the introduced mountain goat (Oreamnos americanus). Both Nuttall's cottontail and the white-tailed jackrabbit are more
abundant on the Limestone Plateau than elsewhere in the Hills. Because of the plateau's broad valley meadows and proximity to the foothills, it also provides optimal habitat for both the mule deer (Odocoileus hemionus) and white-tailed deer (O. virginianus) . Streams transect all of these areas to produce the Un- differentiated Alluvial Soil Association that is inhabited by various riparian spe- cies such as the masked shrew (Sore.t cinereus) and meadow jumping mouse {Zapus liudsonius) .
Faunas of two isolated Tertiary ig- neous protrusions are noteworthy. Inyan Kara Mountain, surrounded by foothills and upland prairie in southeastern Crook County, Wyoming, is the primary range for a herd of recently introduced moun- tain sheep (OvLs canadensis) a species formerly native to the Hills (Baird, 1858: 678; Cowan, 1940:543). Devils Tower is an igneous monolith that proti-udes 865 feet above the surrounding prairie and open pine woodland. The fissures and surface atop the Tower are well carpeted with lichens, grass, sagebrush, and prick- lypear cactus, and support a meager fauna consisting of least chipmunks, deer mice, and bushy-tailed woodrats {Neo- toma cinerea) (exhibit. Visitor's Center, Devils Tower National Monument).
Higher areas in the foothills support a fauna that is similar to that of the boreal-cap, whereas the fauna of the lower regions more closely approaches that of the surrounding prairies. For example, along the foothill-Limestone Plateau border, the meadow vole often is captured in the same trapline with the long-tailed vole. Along the foothill-Red Vallev transition, the meadow vole is associated with the prairie vole {Micro- tus ochrogaster) , and the long-tailed vole is not present. Typical mammals of the Red Valley grasslands and lower foothills are the desert cottontail {Syl- vilagus audubonii), white-tailed jack- rabbit, thirteen-lined ground squiiTcl, black-tailed prairie dog {Cynomys liido- vicianus), deer mouse, western harvest mouse {Reithrodontomys megalotis).
18
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Fig. 4. The Needles, exposed crystalline rock in the Mount Rushmore-Hamey Peak area of the Central Basin. These moss- and lichen-covered granitic ridges thrust upward nearly perpen- dicular to the surrounding forest, and comprise the restricted range of the introduced mountain
goat.
Fig. 5. A broad upland valley on the rolling slopes of the Limestone Plateau, south of Moon. Such areas generally are farmed in oats, legumes and tame grasses, or are grazed, but also
support large herds of deer.
TURNER: MAMMALS OF THE BLACK HILLS
19
ml/'i»»i
t "^F?
Fig. 6. Due to resistence to erosion, high cliffs project as steep walls above canyon floors in more rugged areas of the Limestone Plateau, such as Little Spearfsh Canyon. The above photograph was taken from the rear entrance of a cave, in a cliff above the Timon Camp- ground, that served as a well-used night roost for several kinds of bats (see account of small- footed myotis ) .
Fig. 7. The Red \ alley, locally termed the "racetrack," is a remarkably continuous trough that encircles the Black Hills, and occupies a valley-like position between the footliills and the Dakota Hogback. Large herds of bison, elk, and pronghom, and colonies of prairie dogs inhabit the extensive grasslands comprising the Red Valley portion of Wind Cave National
Park.
20
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
^'^■:-j^£n\'
Fig. 8. The outer border of the Red Valley, and of tlie Black Hills, is formed by a steep- sided, 600-foot sandstone ridge, termed the Dakota Hogback, that slopes gradually outward onto the surrounding semiarid mixed grass prairie. The above break in the ridge (Buffalo Gap) has been eroded away l)y Beaver Creek as it flows out of the Black Hills. Large herds of bison and pronghorn previously migrated between the grasslands of the Red Valley and those of the Northern Great Plains through this stream gap, which also pro\ided access for Jedediah Smith in 1823 and later explorers and adventurers.
Fig. 9. The predominant vegetational component of the Black Hills, especially in the Central Basin, is a montane belt of ponderosa pine. This conifer seems tolerant of xerophytic condi- tions and grows even on exposed rocky ridges. The dark appearance of the foliage of the ponderosa pine, when viewed from a distance, accounts for the Sioux Indian name Paha Sapa,
or "Black Hills."
TURNER: MAMMALS OF THE BLACK HILLS
21
Fig. 10. Moist, narrow valleys in the Limestone Plateau support a dense subalpine belt of white spruce with an associated understory of northern affinities. Luxuriant riparian habitats such as that bordering the Beaver Creek Campground are occupied by masked shrews, meadow
jumping mice, and several kinds of voles.
:^ -*
9^-
■^
•%
. </'
Fig. 11. The Bald Hills is one of three isolated grasslands in the Central Basin situated on ^\■ell-rounded slopes underlain by slates. There is no evidence tliat these grasslands have
ever been forested (Photograph by Barry L. Siler).
22
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
prairie vole, badger ( Taxidea taxiis ) , and coyote (Canis latrans). In addition to those species just mentioned, the Sandhill Regosol Soil Subassociation along the Custer County, South Dakota-Weston County, Wyoming border is inhabited by se\'eral heteromyids. Included among the latter are the hispid pocket mouse (Perognathus Impidiis), oHve-backed pocket mouse (P. fasciatus), and Ord's kangaroo rat {Dipodomys ordii); all are characteristic of rocky or sandy soils.
The outer border of the Red Valley is formed by a 600 foot sandstone rim that is steep-faced within, but slopes gradually outward onto the surrounding Great Plains (Fig. 8). This hogback ranges in e]e\'ation from 3800 to 4900 feet (near Elk Mountain). The outward- sloping cuestas frequently are broken by streams. The inner face of the hogback is of Jurassic age and is composed of green shales and red sandstones of the Sundance Formation and to a lesser ex- tent of the shales, sandstones, and lime- stones of the Morrison Formation. The outer slope is judged as Cretaceous in age and formed of the Skull Creek and Inyan Kara formations, both of which are composites of Dakota and Lakota sand- stones, Fuson shale, and Minnewaste limestone formations. The conspicuous hogback, marking the boundary between the Black Hills and the Northern Great Plains, forms a series of ridges that were an obstacle to early travelers. Beaver Creek has eroded away a break in the hogback, just northwest of Buffalo Gap.
It was through this stream gap that large herds of bison (Bison bison) previously migrated into the grasslands of the Red Valley.
CLIMATE
The climate of the Black Hills is dis- tinct from that of the surrounding, semi- arid Great Plains in being more moist and less subject to extremes of tempera- ture. Because of warm "chinook" winds and frequent sunny skies, the Black Hills are the warmest part of South Dakota in winter. An additional moderating influ- ence is the tendency for hea\y colder air to seek low elevations; thus, Arctic air masses that blanket the plains in colder winter months bypass the higher eleva- tions of the Hills. Conversely, summer temperatures at higher elevations in the Hills are cooler than are those of the surrounding prairies.
Contrasts in climate between Lead, in the north-central part of the Black Hills, and Hot Springs, at the southern periphery, are shown in table 1 and fig- ure 12. Average maximum temperatures for July range from 81-83' F in the Black Hills and 88-92 F on the adjacent plains; axerage minimum temperatures for Janu- ary range from 9-14^ F in the Hills and 4-9° F on the surrounding grasslands. Warm days and cool nights are charac- teristic of the Hills. Air temperatures usually have a daily range from 50-60° F in the shade in summer, but the range may reach 80°, or more, in direct sun- light (Wetmore, 1968:217).
Table 1. — Temperatiue and precipitation data for Hot Springs (southeastern Black Hills) and Lead (north-central Black Hills), 1931-1935 (Hodge, 1960:8).
|
Temperature (°F) |
Precipitation (inches) |
|
|
Climatological station ( with altitude ) |
Highest recorded temperature Highest monthly mean Lowest recorded temperature Lowest monthly mean Annual mean |
Highest monthly mean Lowest monthly mean Mean annual precipitation Mean annual snowfall |
|
Hot Springs (3535 feet) Lead (5245 feet) |
112 75.3 —41 25.1 48.8 (July) (January) 101 69.7 —40 " 24.3 44.9 (July) (January) |
3.04 0.36 16.06 36.4 ( May ) ( December ) 4.09 0.87 23.81 100.1 ( June ) ( February ) |
TURNER: MAMMALS OF THE BLACK HILLS
23
80
60-
H- 40-
z o
z <
LU
^ 20
^
'j^
-f
2
4
MEAN MONTHLY PRECIPITATION (INCHES)
Fig. 12. ( broken
Hydrothermograph for Lead (solid line) in the north-central Black Hills, and Hot Springs line) in the southeastern Black Hills, 1931-1935 (Hodge, 1960:8). Labeled circles ( X ) indicate annual means of both parameters for each climatological station.
The frost-free season is shortest at higher elevations, where brief freezing has been known to occur at any time of summer; however, there are usually only 110 frost-free days in the Black Hills as compared to 130 on the pheripheral prairies.
Average annual precipitation in the Black Hills ranges from 16 inches, in the southern portion, to 28 inches in the northern section, where snow and rain often are formed when prevailing winds are forced abruptly up steep sides of mountains. Mean annual precipitation on the adjacent plains is 14-16 inches. Much of the total precipitation occurs as rain during summer in the drier south- ern portion of the Hills, whereas it is much more c\'enly distributed through- cut the year in the moister northern sec-
tion. Hailstorms generally occur in mid- summer, and lightning usually in late summer.
SOILS
Materials from which soils have de- veloped in the Black Hills include an- cient crystalline rock in the central part of the region and sedimentary rocks ( shale, sandstone, and limestone ) in out- lying areas (Figs. 1 and 3). Local topog- raphy contributes to soil formation by determining drainage; thus, steep slopes have well-drained, thin soils, whereas more level areas have more poorly drained, deep soils.
The major soils of the Black Hills are the Gray Wooded Soils; they are unique to the general region because they de-
24
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
veloped under forest in a relatively hn- mid climate. Soils of the surrounding area developed under grasslands in a climate ranging from moist subhumid to semiarid. The horizons of Gray Wooded Soils of the Black Hills are subdivided as follows: 1) Aon (forest litter). Am (partly decayed litter), and A02 (well decayed litter); these are horizons which are organic and thin (i.e., one-half to two inches in depth). 2) Ai — a mineral hori- zon that is thin or absent. 3) A:> — 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<S70's. Mica, silver, tin, coal, tungsten, and uranium mining flour- ished in later periods. As a consequence, old mine shafts and timnels are available for utilization by bats. Also, the mature ponderosa pine forest, and the numerous deserted farm buildings and mining shanties serve as roosting sites for those species preferring these habitations.
Jones and Genoways ( 1967b: 184- 196) summarized the distributional status of bats in South Dakota on the basis of data accumulated prior to 1967. Subsequently, additional obsei^vations on bats from western South Dakota (Turner and Jones, 1968:444-447) and the Black Hills (Turner and Davis, 1970:360-364) have been published. In the following accounts, pertinent data are summarized or cross-referenced in an effort to avoid repetition. Robert A. Martin, South Da- kota School of Mines and Technology, Rapid City, currently is studying several aspects of bat biology in the Black Hills.
Myotis keenii septentrionalis (Trouessart) Keen's Myotis
[Vcs])ertilio ffryphus] var. septentrionalis Trous- sart, 1897, Catalogus mammalium . . ., fasc. 1, p. 131 (type locality, Halifax, Nova Scotia).
Myotis keenii septentrionalis — Miller and G. M. " Allen, 1928, Bull. U. S. Nat. Mus., 144:105, 25 May.
Keen's myotis is known in the Black Hills from Pennington and Custer coun- ties in South Dakota, and from Weston County, in Wyoming. The species was first reported from the area by Miller and Allen (1928:107) from Elk Mountain, Custer County. Its distribution and sys- tematics subsequently were summarized by Jones and Genoways (1967b: 185).
In the summer of 1968, field work in the Hills area revealed Keen's myotis to
44
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
be common locally (Turner and Davis, 1970:360), although previously this spe- cies had been considered to be rare. Ei^ht males were taken in a mist net set within the entrance to Jewel Cave on 24 July (see account of M. lucifiigus). Eight males and one female were netted at the entrance to Ice Cave on 26 July (see account of M. thysanodes), and five males and one female were captured in nets set over two small ponds near Moon on 30 July. An additional female was netted over one of these ponds on 20 August (see account of L. horealis). Ex- tensive use of mist nets, rather than shooting at dusk, undoubtedly contrib- uted to increased success in capturing this bat. At one place or another, all nine other species of bats in the Hills have been netted in company with Keen's my Otis.
The status of M. keenii in the Black Hills in winter months remains uncertain. Specimens have been taken in the area from 9 June through 25 August. R. A. Martin (pers. com.) investigated more than 100 caves and mine shafts in Law- rence, Pennington, and Custer counties in the winter of 1969-70, but M. keenii was not observed in any of these hiber- nacula. Presumably the species repre- sents a disjunct population in the Black Hills region; the nearest localities of record are 252 miles to the north at Fort Buford, Williams Co., North Dakota (Miller and Allen, 1928:106-107), and 296 miles to the southeast at a locality 3 mi SW Springfield, Bon Homme Co., South Dakota (Jones and Genoways, 1967b: 185). Surely, if this species of bat were a transient, it would be in evi- dence along routes of migration. Thus, based on distributional data, the species is assumed to be resident in the Hills throughout the year.
Ecological information pertinent to M. keenii in the Black Hills is limited. Known altitudinal range is between 4000 and 6500 feet. Three specimens collected on 19 July 1951 near Buckhorn, Weston Co., ^^^'()ming, were found roosting in old buildings of a sawmill. A foraging
male was shot about 8:30 P.M. (MDT) on 9 June 1965 over Porcupine Draw; this individual was found with two M. hicifugiis, one M. leihii, 11 M. volans, and one Lasiunis cinereus. Procupine Draw enters Ditch Creek Valley from the east; its steep rocky hillsides are clad in ponderosa pine and white spruce.
A single, lactating female was ob- tained in Ice Cave on 26 July; two other lactating females were taken near Moon on 30 July and 20 August. A subadult male (evidenced by unfused phalangeal epiphyses) was captured at Ice Cave on 26 July. The testes of a male shot in early June were 2 in length, whereas those of 21 males collected in late July had an average length of 4.8 (3-6). Molt was underway on 10 males taken in late July, but not on 11 other speci- mens taken in the same period. A fe- male obtained on 20 August was in complete new pelage. The ears of the male shot in Porcupine Draw were in- fested by chiggers, Leptotromhidium myotis ( E wing ) .
The taxonomic status of Keen's my- otis in the Black Hills merits further in- vestigation. I concur with the opinion of Jones and Genoways ( 1967b: 185-186) that individuals from the Hills fall within the currently understood range of vari- ation of M. k. septentrionalis from Kan- sas, Nebraska, Iowa, and eastern South Dakota (Tables 6 and 7). Although there are no significant sexual differences among Black Hill specimens, females are slightly larger than males in all ex- ternal and cranial measurements. The pelage of females is also slightly paler in tone of color and more intense in re- flectance of reds and blues; whereas the pelage of males reflects greens more in- tensely. The average breadth of the zygomatic arch, braincase, upper molars, and mastoid (excluding specimens from Iowa and Kansas) is narrower in indi- \'iduals from the Hills than in specimens from areas to the east and southeast. Moreover the forearms of the specimens from the Hills are slightly shorter (ex- cluding those from eastern South Da-
TURNER: MAMMALS OF THE BLACK HILLS
45
Table 6. — Geographic variation in selected external and cranial measurements of adult Myotis
keenii from the central United States.
|
Number and se.x of specimens averaged |
Total length |
Forearm length |
Zygomatic breadth Breadth of braincase |
Breadth across upper molars |
Mastoid breadth |
|
37 (3U,69) Mean |
89.0 |
Black Hills, South Dakota and Wyoming 34.9 8.9 7.1 5.5 |
7.6 |
||
|
SD |
±3.49 |
±1.06 |
±0.20 ±0.25 |
±0.15 |
±0.22 |
|
4 (2c^,29) Mean |
87.0 |
Bon Homme County, South Dak 34.7 9.3 7.3 |
ota 5.7 |
7.9 |
|
|
SD |
±3.37 |
±0.77 |
±0.22 ±0.08 |
±0.03 |
±0.16 |
|
12 (5c?, 79) Mean |
92.5 |
35.4 |
Sarpy County, Nebraska 9.3 7.5 |
5.7 |
7.7 |
|
SD |
±5.18 |
±0.76 |
±0..38 ±0.32 |
±0.18 |
±0.24 |
|
11 (35,89) |
Boone, |
Hardin, and Keokuk counties, Iowa |
|||
|
Mean |
88.6 |
36.1 |
9.1 7.2 |
5.6 |
7.5 |
|
SD |
±4.63 |
±0.95 |
±0.20 ±0.16 |
±0.20 |
±0.17 |
|
12(76,59) Mean |
87.1 |
.3.5.1 |
Marshall Countv, Kansas 9.1 7.4 |
5.7 |
7.5 |
|
SD |
±3.89 |
±0.82 |
±0.22 ±0.25 |
±0.11 |
±0.18 |
kota), and their total body length is greater (excluding those from Ne- braska). Additional measurements (not listed in Table 6) of 37 specimens from tlie Black Hills are: tail length, 39.4± 2.57; hind foot length, 9.5±0.56; ear length, 17.2 ±0.90; greatest length of skull 14.9 ±0.29; and maxillary tooth- row length, 5.8 ±0.17.
There are no significant geographic differences in the color of pelage of indi- viduals collected in the summer; how- ever. Keen's myotis from the Black Hills are darker on the average than those from Nebraska, Kansas, and eastern South Dakota. Reflectance of hues also varies geographically (Table 7). Specimens from the known population nearest the Hills (Bon Homme County, South Da- kota) reflect reds and blues less in- tensely, and greens more intensely than representatives from the Hills.
Specimens examined (42). — SOUTH DA- KOTA: Pennington County: Moon, 22 mi W Hill Cit>', 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<i,ans) . The bat was wedged in a horizontal crevice wherein the ambient temperature was 7.2° C. Four solitary males taken from Jewel Cave on 20 November 1967 had rectal temperatures of 6.4°, 7.1°, 7.5°, and 8.0°C; corresponding ambient temperatures were 5.6°, 5.6°, 7.2°, and 6.7°C. Approximately 600 P. townsendii were in Jewel Cave at this time. On 7
February 1970, four male M. leihii were found hibernating in French Creek Cave located in Custer State Park (see ac- count of E. fuscus ) .
Small-footed myotis taken in the course of this study usually were found near a source of water. Two males and three females were shot in mid-June as they foraged over a stock pond that formed the dammed upper portion of a pine-filled draw near Minnckahta. A male was shot in early July as it flew over Boxelder Creek, near Nemo, and another was shot in late July as it flew over the Savoy Reservoir in Little Spearfish Can- yon. Two males were netted over a woodland pond near Moon on 30 July 1968 (see account of L. horealis), and another was shot over a reservoir at Roby Springs on 6 September. At several lo- calities, M. lucifugus, M. keenii, M. thy- sanodes, M. volans, L. noctiva<^ans, E. fuscus, and Lasiurus cinereus have been shot or netted as they foraged in the same areas as M. leihii.
This species retires to night roosts after feeding, as do most other kinds of vespertilionids. All specimens from the
TURNER: MAMMALS OF THE BLACK HILLS
47
Cambria Coal Mine were entering the shaft when netted between 21:15 and 22:25 hrs (MDT). While collecting in Little Spearfish Canyon in late July and early August 1967, I located a well-used night roost abox'c Tinion Campground. One entrance to the cave was about three feet in diameter, but a second entrance, 10 by 15 feet, opened out over the sheer 200-foot cliff. Most of the cave was open, roomy, and well lighted; however, to tlie rear, an elongate, narrow chimney rose about 40 feet above the ceiling. Although the cave often was visited dur- ing the day, only one bat (a male P. townsendii) was taken; nevertheless, at night several bats were seen clinging to the walls of the chimney while many others flew in and out of the large en- trance overlooking the cliff. One male M. leihii, five M. hicifugus, and 10 M, volons were collected in the roost at this time (Turner and Jones, 1968:444). On 30 July 1968, Wayne H. Davis and I returned to the roost. We spent the night within the cave and established a net angling across the cavern; two additional male M. leihii were taken, along with 24 M. hicifugus and 42 M. volans (Turner and Davis, 1970:361).
Although widespread throughout the Hills, populations of the small-footed myotis seemingly are relatively small. However, on 24 July 1968, Davis, Roger Barbour, and I captured 48 individuals (five females and 43 males) in a net set back within the entrance of Jewel Cave; 35 of these were banded and released (see account of M. hicifugus). Thus, small-footed myotis may congregate at favorable sites in the Black Hills.
Females of this saxicolous species usually bear only a single young. Jones and Genoways (1967b: 187) reported that no embryos were found in the South Dakotan specimens examined by them and that most individuals collected in the summer were males. Of three fe- males obtained near Minnekahta in mid- June 1967, one contained an embryo that was 8 in crown-rump length. A female taken northeast of Rapid City on 17
July 1945 carried an embryo that was 13. Testicular lengths of 17 males obtained from mid-June to late July were 3 or 4, whereas those of three November-taken males were 2. The two males from Moon and many of the specimens from Jewel Cave were subadults in conspicuous molt. Small-footed myotis from the Black Hills are parasitized by a chigger, Lepto- twmbiclium myotis (Ewing), and a fe- male from Jewel Cave harbored a bat bug, Cimex piloseUus (Horvath).
Specimens examined (66). — SOUTH DA- KOTA: Lawrence County: Cleopatra Mine, 1.5 mi N Carbonate, 2 (SDMT); Little Spear- fish Canyon, Savoy, 8 nii W Lead, 5200 ft, 1; Little Spearfish Canyon, 2 nii S, 10 mi W Lead, 6000 ft, 3; 2 mi W Nemo, 4700 ft, 1. Meade Cuuntij: Davenport Cave, 3 mi S, 0.5 mi W Sturgis,' 4400 ft, 5 (NRW). Pennington County: 8 mi ENE Rapid City, 1 (UMMZ); Rapid City, 1 (SDMT); Dark Canyon, 2 mi S, 5 mi W Rapid City, 3800 ft, 1; Diamond S Ranch, near Rapid City, 1 (UMMZ); Moon, 22 mi W Hill City, 6200 ft, 2; Porcupine Draw, 14 mi W Hill City, 6400 ft, 1; Igloo Cave, 3 mi S, 16 mi W Hill City, 2 (SDMT); T. 2 S, R. 6 E, E ]2 sec. 19, 1 (MSWB). Custer County: 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; Roby Springs, 4 mi N, 22 mi W Custer, 5400 ft, 1; Hell's Canyon, 13 mi W Custer, 3 (USNM); Custer, 1 (AMNH); French Creek Cave, Custer State Park, 4 (SDMT); Jewel Cave, 2.5 mi S, 12 mi W Custer, 5280 ft, 20 (1 NRW, 1 SDMT, 4 UK, 1 UW); S and G Cave, 11 mi SW Custer, 1 (SDMT). Fall River County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 5.
WYOMING: Weston County: Cambria Coal Mine, 6 mi N Newcastle, 6000 ft, 8.
Additional record.— SOUTH DAKOTA: Custer County: Wind Cave National Park (Peck, 1964:38).
Myotis lueifugus carissima Thomas
Little Brown Myotis
Myotis (Leuconoe) carissima Thomas, 1904, Ann. Mag. Nat. Hist., ser. 7, 13:383, May ( type locality, Yellowstone National Park, Wyoming ) .
Myotis lueifugus carissima — Gary, 1917, N. Amer. Fauna, 42:43, 3 October.
Myotis hicifugus is distributed throughout the northern half of North America and evidently widespread and fairly common in the Black Hills. The known altitudinal distribution of this species extends up to 6900 feet. With
48
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
the exception of three individuals ob- tained in mid-x\ugust and one taken in early September, all specimens examined were collected in June and July. Only two females (from Crook County, Wy- oming) taken on 1 July 1947 and 24 August, 1913, were among the specimens studied, although Jewel Cave has been reported as a hibernaculum for both sexes (Jones and Genoways, 1967b: 187).
In the summer, females ordinarily segregate in nursery colonies; this may account for their scarcity at this time. On 4 July 1970, Robert A. Martin (pers. com.) set a mist net between two chim- neys in the attic of the Bob Marshall Camp dining hall, located on the shores of Bismark Lake, about 1 mi E Custer. In the morning, the net yielded seven adult females, one subadult male, and a newly born young. Martin estimated that 100 to 150 adults comprised the nursery colony. To my knowledge, this is the first report of a maternity colony of M. lucifugus in the Black Hills region. Martin also captured two females in Rapid City in the summer of 1970.
Mijotis lucifugus carissima contrasts with M. I. lucifugus of eastern South Dakota in being conspicuously more pale and in having larger average external and cranial dimensions (Jones, 1964 :(S6 and Table 5). The subspecific name carissima was assigned to specimens from the Wy- oming portion of the study area by Miller and Allen (1928:52), and later to speci- mens from South Dakota by Jones and Genoways (1967b: 187).
Many little brown myotis collected in the course of this study were collected (shot or netted) while the bats were foraging over water, grassy meadows or along the edge of stands of ponderosa pine or white spruce; several individuals were netted at cave entrances as well. Some typical situations in which M. lucifugus were obtained are as follow: in early July 1965 seven individuals were collected, along with two M. volans, over a beaver pond near Beaver Creek Camp- ground, where the greatest activity oc- curred just after dark: two were shot on 3 Jul)' 1967 over a stock pond in a
bluegrass meadow 1 mi E Nemo along with one Lasiuius cinereus, one Lasion- ycteris noctivigans, and one Eptesicus fuscus; one was netted on the same date over Boxelder Creek, in Lawrence County along with seven L. cinereus and one E. fuscus; one was shot as it flew over a reservoir at Roby Springs on 6 September 1968; two more were netted over a \\'oodland pond at Moon, one each on 30 July and 20 August 1968 ( see account of L. borealis); another was netted on 26 July 1968 at the entrance of Ice Cave (see account of M. thy- sa nodes).
On 13 June 1965, members of a field party from the Museum of Natural His- tory obtained 12 M. lucifugus, five M. volans and a mummified Plecotus town- semlii from Jewel Cave. The majority of the Myotis were located in the "dungeon" portion that was wet with seepage water and had an ambient tem- perature of 6.7°C; all were torpid. Five M. lucifugus, 48 M. leibii, eight M. keenii, 10 M. volans, one E. fuscus, and four P. townsenclii were netted at the entrance to Jewel Cave on 24 July 1968; most of the bats were entering the cav- ern at the time of capture, between dusk and midnight. Fi\'e little brown myotis were caught at a night roost in Little Spearfish Canyon in late July of 1967 (see account of M. leibii), and another 24 (of which 14 were banded and re- leased) were netted there on 30 July 1968.
The average testicular length of 25 males taken in June and July was 4.2 (2-7). A female, with a young attached, was extracted from a crack in the wall of a summer cottage 15 mi ENE Sundance, Wyoming, on 1 July 1947.
Molt was apparent in an adult male taken on 7 July 1965, in Pennington County, because new pelage was ob- served underlying the old in the mid- dorsal region and behind the ears. Sev- eral individuals from Little Spearfish Canyon, obtained on 30 July, were sub- adults in conspicuous molt. An albinis- tic little brown bat was captured in Jewel Cave on 31 August 1966 by L. N. Brown
TURNER: MAMMALS OF THE BLACK HILLS
49
and associates of tlie University of Wy- oming.
A male taken at Moon on 20 August 1968 was parasitized externally by bat flies, Bosilia forcipata Ferris, and by lae- laptid mites, Macronyssus crosbtji (Ewing and Storer). Mijotis lucifugus caiissima from many localities in the Hills harbored chiggers, Leptotrom- bidiinn myotis (Ewing). In a report filed at Wind Cave National Park, Olaf E. R\'berg recorded another species of bat fl)', Trichobiiis corynorJiiniis Cockerell, on specimens taken at Jewel Cave in May 1962.
Spc'citncns examined (84). — SOUTH DA- KOTA: Lawrence County: Little Spearfish Canyon, Savoy, 8 mi W Lead, 5200 ft, 3; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 6000 ft, 15; 2 mi W Nemo, 4700 ft, 4; 3 mi E Nemo, 4700 ft, 1; 1 mi E Nemo, 4700 ft, 3. Meade Coiiniy: Haven Dams, 6 mi S, 1.5 mi W Stui- gis, 4600 ft, 1. Pennington County: Rapid City, 1 (NRW); Sitting Bull Cave, 1 (SDMT); Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 8; 3 mi N, 7 mi W Deerfield, 6900 ft, 1; Moon, 22 mi W Hill City, 6200 ft, 2 (1 UK); Ditch Creek, 14 mi W Hill City, 6400 ft, 2; T. 2 S, R. 6 E, E M sec. 19, 1 (MSWB); 10.5 mi E Wyoming border, 5.8 mi N Pennington-Custer county line, 6 (MSWB). Custer County: 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; Roby Springs, 4 mi N, 22 mi W Custer, 5400 ft, 1; Jewel Cave, 2.5 mi S, 12 mi W Custer, 5280 ft, 27 (2 SDMT, 8 UW, 5 UK); Housing Area, Wind Cave National Park, 4100 ft, 2 (WCNP). Fall River County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 1.
WYOMING: Crook County: Sand Creek, 3750 ft, 2 (USNM); 15 mi ENE Sundance, 3825 ft, 1.
Additional records ( see te.xt also ) . — SOUTH DAKOTA: Lawrence County: Spearfish (BB).
Myotis thysanodes pahasapensis
Jones and Genoways
Fringe-tailed Myotis
Myotis thysanodes pahasapensis Jones and Genoways, 1967, Jour. Mamm., 48:231-235, 20 May ( type locality, 6 mi N Newcastle, 6000 ft, Weston Co., Wyoming).
On 2 July 1965, I removed a fringe- tailed myotis from a net stretched across a horizontal shaft in the deserted Cam- bria Coal Mine complex, 6 mi N New- castle, Weston Co., Wyoming. This male subsequently was designated as the holo- type of Myotis thysanodes pahasapensis,
a subspecies restricted to the Black Hills and adjacent areas. The range of this subspecies is disjunct from the more western range of M. t. thysanodes by approximatelv 400 miles (Jones and Genoways, i967a:234; 1967b: 188). In South Dakota, Myotis thysanodes has been taken only from Pennington, Custer, and Fall River counties, and in Wyoming only from Weston County. Its known altitudinal range in the Hills is between appro.ximately 3800 and 6200 feet.
Thirty years intervened between the first report (Bole, 1935:147-148) and the second report (Thompson, 1965:37) of this species in the Hills region. In the interim, a young female from Custer State Park (Stebler, 1939:389) and an adult male from 1.5 mi E Buckhorn, Wy- oming (Long, 1965:532) were reported as Myotis evotis.
The fringe-tailed myotis resides in the Black Hills the year around, as evi- denced by a hibernating male taken in a cave along Boxelder Creek near Dody Spring, in Pennington County, on 17 February 1968 (Turner and Jones, 1968: 445). Methods of capture of specimens collected previously are unknown. Indi- viduals obtained in the present study either were shot or netted as they for- aged in the following areas: 1) over water — one female (5 July 1966) over a stream 3 mi N Hot Springs; one female (13 July 1967) over a stock pond in a pine-filled draw near Minnekahta; one male (30 July 1968) and two females (30 July and 20 August 1968, respec- tively) over a woodland pond near Moon (see account of L. borealis; 2) over a grassy meadow — a male (19 July 1951) feeding at dusk 1.5 mi E Buckhorn in company with one Eptesicus fiisciis; 3) at the entrances of caves — one female and five males (July and August, 1965- 66) and one male (17 February 1970) at Jewel Cave (see account of M. hici- fugtis); two males (26 July 1968) at Ice Cave (see account of E. ftiscus); 4) at the opening of a mine shaft — the holo- type of pahasapensis (taken along with M. lei])ii). In addition, three feniah^s
50
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
were taken in mid-June 1968 while in a day roost, which was situated among tlie rafters of an open porch in Wind Cave National Park; these fringe-tailed myotis were clustered in close association with M. volans (Turner and Davis, 1970: 361).
When Wayne H. Davis and I netted the entrances to Ice Cave on 26 July 1968, a flurry of activity occurred be- tween 21 and 22 hrs ( MDT), and gradu- ally decreased thereafter. Nine M. keenii, one M. lucifugus, 20 M. volans, and four E. fuscus were captured with M. thy- sanodes. The large mass of natural ice that persists in the cavern, even in sum- mer, is indicative of the cool tempera- tures therein.
Each of the three females obtained at Wind Cave National Park on 15 June carried a single embryo; they were 6, 8, and 12 in crown-rump length. Lac- tating females were taken at Moon on 30 July and 20 August. A male also cap- tured on 30 July had testes that were 5 in length. Young of the year were noted flying as early as 10 August.
New hairs were found beneath the old pelage of males taken in late July. Molt had not begun on a female taken at the same time, but it was relatively far advanced on a female obtained on 20 August. Three pregnant females cap- tured at Wind Cave National Park on 15 June were molting over both dorsum and venter.
A female fringe-tailed myotis from Fall River County harbored a bat fly, Basilia forcipata Ferris, chiggers, Lepto- trombicUiim myotis (Ewing), and a bat bug, Cimex adjunctus Barber. Dr. George W. Byers, Department of Entomology, The University of Kansas, identified the remains of a moth, a small scarabid beetle, and a large alleculid beetle from the stomach of this same individual.
In contrast with Myotis evotis, which the fringe-tailed bat resembles superfi- cially, Myotis thysanodes possesses a longer forearm, shorter ears, slightly larger cranial dimensions, a conspicuous fringe of hairs on the free border of the uropatagium, a better developed sagittal
crest, a slightly higher and more inflated braincase, and somewhat more robust teeth (see Jones and Genoways, 1967a: 233-234; 1967b: 189). Although M. evo- tis occurs over much of Wyoming and western South Dakota, no specimens have been obtained in the Black Hills. Jones and Genoways (loc. cit.) sug- gested that thysanodes and evotis might be ecologically separated.
Fringe-tailed myotis from the Black Hills differ from M. t. thysanodes in having larger ears, shorter forearms, more contrast in color between dorsal pelage and membranes, and a smaller skull that is relatively narrow ( Jones and Genoways, 1967a: 233). After prelimi- nary analysis for secondary sexual vari- ation, the pelage of individuals collected in the summer from east-central Cali- fornia (Tuolumne County) and from Cocomino, Cochise, and Gila counties, Arizona, was compared with that of rep- resentatives from the Black Hills. Cali- fornian specimens are significantly darker in color tone than those from either Ari- zona or the Black Hills. Major differ- ences between the latter two populations are the greater reflection of greens by representatives from the Hills and the slightly lighter tone of the individuals from Arizona. Average measurements and standard deviations of pelage color of 10 ( four males and six females ) adult M. t. pahasapensis collected in the sum- mer are: color tone, 25.3 ±4.22; percent red reflectance, 55.0 ±0.56; percent green reflectance, 20.1 ±0.29; and percent blue reflectance, 24.9 ±0.36. Selected average measurements and standard deviations of 16 specimens, nine males and seven fe- males, of fringe-tailed myotis from the Black Hills are: total length, 94.4 ±3.22; tail length, 41.0±1.94; hind foot length, 10.6±0'!^92; ear length, 18.5±0.95; fore- arm length, 41. 1± 1.20; greatest length of skull, 16.6 ±0.27; zygomatic breadth, 10.1 ±0.21; interorbital constriction, 3.9 ±0.14; braincase breadth, 7.7 ±0.19; mastoid breadth, 8.3 ±0.19; breadth across upper molars, 6.5±0.22; maxillary tooth-row length, 6.3 ±0.16; and breadth across upper canines, 3.9±0.18.
TURNER: MAMMALS OF THE BLACK HILLS
51
S))ccimens examined (30). — SOUTH DA- KOTA: Pennington County: along Boxclder Creek, near Dodv Spring, 4060 ft, 1; Moon, 22 mi W Hill Cit\, 6200 ft, 3; Ice Cave, 8 mi N, 15 mi W Custer, 6400 ft, 2 (UK); 10.5 mi E \\'\oming border, 5.8 mi N Pennington- Custer count)- line, 7 (MSWB). Custer County: Grace Coolidge Creek, Custer State Park, 1 (UMMZ); jewel Cave, 2.5 mi S, 12 mi W Custer, 5280 ft, 9 (3 SDMT, 1 UMMZ, 4 UW); Ele\'ator Building, Wind Cave National Park, 4100 ft, 3; 3 mi N Hot Springs, 1 (UW). Fall River County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 1.
\\TOMI\G: Weston County: 1.5 mi E Buckliorn, 6150 ft, 1; Cambria Coal Mine, 6 mi N Newcastle, 6000 ft, 1.
Myotis volans interior Miller Long-legged Myotis
Myotis longicrus interior Miller, 1914, Proc. Biol. Soc. Washington, 27:211, 31 October (type locality, 5 mi S Twining, 11,300 ft, Taos Co., New Me.xico).
Myotis volans interior — Miller and G. M. Allen, 1928, Bull. U. S. Nat. Mus., 144:142, 25 May.
Myotis volans, first reported in the study area by Moulthrop (1936:413), is known throughout the Black Hills, ex- cept from Crook County, Wyoming. Evi- dently this is the most common and widely distributed species of the genus in the region. In general, this species occupies montane areas of western North America. The known altitudinal range in the Black Hills is 4000 to 6500 feet, but the species is most abundant above 4500 feet. It is resident throughout the year as evidenced by hibernating indi- viduals taken from Bush's Cave (20 No- vember) and Jewel Cave (20 to 26 No- vember and 17 February).
The long-legged myotis superficially resembles the little brown myotis, but M. volans differs from M. hiciftigus in having a shorter rostrum, a braincase that is abruptly elevated from the rostral level, low rounded ears, a keeled calcar, and pelage that usually extends onto the underside of the wing to a line joining the elbow and knee.
Myotis volans forages over such areas as campgrounds, meadows, and water courses. For example, in early June 1965, 11 individuals were shot at dusk as they
flew through Ditch Creek Valley and associated Porcupine Draw in Penning- ton County (see account of M. keenii). Five males and two females were taken in company with one M. leihii over a reservoir at Roby Springs, Custer County, in early September 1968. Five males and two females (four of these bats were banded and released) were netted over ponds near Moon in late July and mid- August of 1968 (see ac- count of L. horealis). A female, evi- dently disturbed from her daytime roost, was captured in mid-morning (10:30 A.M., MDT) at the Beaver Creek Camp- ground, Pennington County, on 13 June 1965. A male collected at this same site on 2 July 1965 carried the remains of a crane fly (Tipulidae) in his mouth.
Caves are utilized in summer by M. volans both as daytime retreats and as nighttime roosts following foraging flights. Individuals have been obtained during the summer from Bear Trap Cave, Ice Cave ( 19 banded and released on 24 July 1968), and Jewel Cave. (Five tor- pid bats were obtained in the latter cav- ern on 13 June 1965; see account of M. liicifugus. Nine M. volans were banded and released in Jewel Cave on 24 July 1968.) On 5 July 1967, I shot a male about 10:30 P.M. (MDT) as it flew into Davenport Cave and alighted on a crys- talline calcite (dog-tooth spar) wall. Ten males were collected from a night roost in Little Spearfish Canyon between 26 July and 7 August 1967 (see account of M. leihii), and 42 males were netted, banded, and released in this same cavern on 30 July 1968. All chiropteran species known to occur in the Black Hills were associated with M. volans at some local- ity. On two different occasions, individ- uals were located in a day roost on Wind Cave National Park in company with M. thysanodes. Eleven of these were banded and released, and a female was netted one month later within 200 yards of the original site of capture (Turner and Davis, 1970:362).
On 20 November 1967, a single male was taken in Bush's Cave, a small cave several hundred feet south of the en-
52
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
trance to Jewel Cave. The ambient tem- perature was 9.7 °C and the rectal tem- perature of the bat, which was torpid and covered by droplets of moisture, was 10.6° C. A single Plecotns townsendii also was noted in Bush's Cave at that time.
Jones and Genoways ( 1967b : 190 ) reported two females, each containing a single embryo, that were obtained on 29 and 30 June in Harding County, north of the Hills. However, of 18 females collected in the summer and examined from the Hills region, none was lactat- ing and only one was pregnant. The latter specimen was taken in Wind Cave National Park on 25 July and the single embryo was 16 in crown-rump length. A male flying young of the year was ob- tained at Roby Springs on 7 September. Testes of 14 males taken from mid-June to late July were 1 to 5 in length.
New hairs were manifest under the old pelage over much of the body of a male taken on 30 July 1961. Bat flies, Basilia forcipata Ferris, were found on M. volans from Pennington, Custer and Fall River counties, whereas other indi- viduals from Pennington, Fall River, and Lawrence counties harbored chiggers, Leptotrombidium mijotis (Ewing). Two females from Wind Cave National Park were infested by spinturnicid mites, Spinturnix americanus (Banks), and a male from Roby Springs, Custer County, was parasitized externally by several un- identifiable (badly damaged) mites.
Specimens examined (77). — SOUTH DA- KOTA: Lawrence County. 2 mi S Tinton, 6100 ft, 1; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 6000 ft, 10; 2 mi W Nemo, 4700 ft, 1. Meade County: Davenport Cave, 3 mi S, 0.5 mi W Sturgis, 4400 ft, 1. Pennington County: Rapid City, 3 (NRW); Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 3; Moon, 22 mi W Hill City, 6200 ft, 3; Ditch Creek, 14 mi W Hill City, 6400 ft, 11; Ice Cave, 8 mi N, 15 mi W Custer, 6400 ft, 1; Bear Trap Cave, 4 mi S, 16 mi W Hill City, 6000 ft, 1; T. 2 S, R. 6 E, E M sec. 19, 1 (MSWB); 10.5 mi E Wyoming border, 5.8 mi N Custer County line, 8 (MSWB); unspecified locality, 1 (NRW). Custer County: Roby Springs, 4 mi N, 22 mi W Custer, 5400 ft, 7; Bull Springs, 2 mi N, 9 mi W Custer, 6500 ft, 1 (UMMZ);
Jewel Cave, 2.5 mi S, 12 mi W Custer, 5280 ft, 12 (3 NRW, 1 SDMT, 1 UW); Headquar- ters, Wind Cave National Park, 4100 ft, 1 (UMMZ); Elevator Building, Wind Cave Na- tional Park, 4100 ft, 1; Housing Area, Wind Cave National Park, 4100 ft, 5. Fall River County: 5.5 mi E Minnekahta, 4000 ft, 1; 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 2.
WYOMING: Weston County: 1.5 mi E Buckliorn, 6150 ft, 1; Cambria Coal Mine, 6 mi N Newcastle, 6000 ft, 1.
Additional record.— SOUTH DAKOTA: Custer County: Wind Cave, Wind Cave Na- tional Park (Peck, 1964:39).
Lasionycteris noctivagans (Le Conte) Skiver- HAIRED Bat
V[espertilio]. noctivagans Le Conte, 1831, in McMurtrie, The animal kingdom ... by the Baron Cuvier . . ., 1:431 (type local- ity, eastern United States).
[Lasionycteris] noctivagans — Peters, 1866, Mo- natsb. k. preuss. Acad. Wiss., Berlin, p. 648 (for 1865).
The silver-haired bat occurs through- out the United States, but is known in the Black Hills from only seven localities in South Dakota; all but two specimens were obtained in transitional areas be- tween the high plains and foothills along the southern and eastern periphery of the Hills. This species first was reported in the study area by Jones and Genoways (1967b: 190). Although little is known of the actual migratory routes, Lasion- ycteris presumably migrates through the Hills region (northward in spring and southward in late summer and early autumn). Jones and Cenoways (loc. cit. ) suggested that this species probably is resident in the higher part of the Black Hills in summer. The altitudes at which most indi\iduals ha^■e been collected thus far generally range from 3800 to 4700 feet, but two males were obtained at 6200 feet near Moon on 29 July, 1968 (see account of L. horealis). Thus, the silver-haired bat is more widely distrib- uted altitudinally than previously sup- posed.
There are few records of silver-haired bats occurring in caves. A solitaiy male was taken from an unnamed cave in Dark Canyon on 19 November 1967 in a semi- torpid state. The limestone ca\'e is lo-
TURNER: MAMMALS OF THE BLACK HILLS
53
cated about 90 feet below the crest of a 200-foot cliff. Relative humidity within the cave was 57 percent, and the ambient temperature was 6.7 ""C; rectal tempera- true of the bat was 9.6^C. The male was hidden in a horizontal crevice and be- came somewhat acti\'e shortly after being extracted. Three Plecotus townsendii and one Myotis leibii, all in a torpid state, also were collected in the same cavern at this time. The lateness of the season, the low rectal temperature, and tlie situation in which the bat was found, all suggest that this individual was not a late transient, but rather in hibernation (Turner and Jones, 1968:445). Thus, it is probable tliat at least a few silver- haired bats reside in the Black Hills throughout the entire year.
In 1967, five individuals, two from southwest of Minnekahta (12 and 13 June) and three from near Nemo (29 June to 3 July ) , were shot as they foraged in early evening in company with Myotis leibii, M. hicifugtis, M. thysanodes, Ep- tesicus fuscus, and Lasiurus cinereus. Another individual was captured at the fonner site at 13:30 (MDT) on 14 June in a net over a small algae-covered pond, and a male was captured in a net set over a sewage settling pond at the head of Wind Cave Canyon on 25 July 1968. This species seems to prefer to forage in grassv vallevs that contain a source of standing water and which are sur- rounded by hillsides well-forested with p'jnderosa pine.
Females give birth to one or usually two young between late May and early July. A female shot near Minnekahta on 12 June 1967 carried two embryos that were 11 in crown-rump length. Length of the testes of five males taken from 13 June to 3 July 1967 varied be- tween 4 and 5; a male obtained in No- vember had a testicular length of 4. The testes of three males obtained in late July 1968 were 6, 7, and 7 in length. Fine silver-tipped hairs were seen underlying the old pelage on the dorsum of the three latter individuals.
A male from Minnekahta, Fall River
County, was parasitized externally by chiggers, Leptotrojnhidium myotis (Ewing), whereas the male from Dark Canyon harbored laelaptid mites, Mac- ronyssus tinidens Radovsky. In addition, B. V. Peterson, Entomology Research Institute, Ottawa, Ontario, identified a bat fly found on a male L. noctivaiians from Wind Cave National Park as either Trichohitis adamsi Auguston or a new species; T. adamsi has been reported previously only from Macrotus californi- cus and Tadarida hrasiliensis.
Specimens examined (11). — SOUTH DA- KOTA: Lawrence County: 2 mi W Nemo, 4700 ft, 2; 1 mi E Nemo, 4700 ft, 1. Penning- ton County: Diamond S Ranch, near Rapid City, 1 (UMMZ); Dark Canyon, 2 mi S, 5 mi W Rapid City, 3800 ft, 1; Moon, 22 mi W Hill City, 6200 ft, 2. Custer County: Wind Cave Canyon, Wind Cave National Park, 4100 ft, 1. Fall River County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 3.
Eptesicus fuscus pallidus Young Big Brown Bat
Eptesicus pallidus Young, 1908, Proc. Acad. Nat. Sci. Philadelphia, 60:408, 14 October (type locality, Boulder, Boulder Co., Colo- rado ) .
Eptesicus fuscus pallidus — Miller, 1912, Bull. U. S. Nat. Mus., 79:62, 31 December.
The pallid big brown bat, originally reported from the region as Adelonyc- teris fusca (J. A. Allen, 1895a:273), is common, widespread, and occurs in all counties that comprise the Black Hills. This species inhabits most of North America and is known in the study area from elevations between 3500 and 6200 feet. Eptesicus fuscus pallidus differs from E. f. fuscus which is geographically adjacent to the east in South Dakota in its discernibly paler pelage, and smaller average external and cranial dimensions (Jones, 1964:92).
Most of the specimens examined were collected in summer; however, individ- uals have been obtained in the winter at Custer (23 January 1888) by Vernon Bailey, in Hell's Canyon (5 January 1902) by Merritt Cary, and in hibemac- ula such as Jewel Cave (Jones and Geno- ways, 1967b: 192). On 7 February 1970,
54
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Robert A. Martin and associates (pers. com.) located a winter colony of 20 E. fusctis, four Myotis leibii, and two Pleco- tus townsenclii in a cave along French Creek in Custer State Park; all bats found in the cavern at that time were males.
While attempting to obtain flying squirrels in a spruce-filled canyon on 11 February 1946, J. A. King captured a female big brown bat by the wing in a rat trap; the trap was attached about seven feet above the ground to a tree, and peanuts were used as bait. The local conditions under which this unseasonal activity occurred are unknown to me. King later collected a male (4 March 1946) and a female (21 March 1946) that were hibernating in an old hori- zontal mine shaft in the same area ( 3 mi SE Hill City, Pennington County).
Many of the big brown bats collected in this study were shot at dusk as they foraged over grassy, conifer-bordered meadows, or over ponds and water courses (Turner and Jones, 1968:446). Others were captured in mist nets in the following situations: One male (3 July 1967) captured as it flew along Boxelder Creek, 2 mi W Nemo (see account of M. lucifu<i,us) . Four males (29 July 1968; three banded and released) netted while foraging over a woodland pond near Moon (see account of L. horealis). One male (24 July 1968) taken at the en- trance to Jewel Cave (see account of Myotis lucifiigus). Four males and one female (26 July 1968; four banded and released) netted at the entrance of Ice Cave. The main activity of 32 bats taken in the last-mentioned cavern was be- tween 21 and 22 hrs (MDT); however, big brown bats were not obtained until about 23 hrs (see account of Myotis thysanodes) . Other species taken along with Eptesicus at various sites of captiuc^ were Myotis keenii, M. leihii, M. hici- fugus, M. thysanodes, M. volans, Lasi- onycteris noctivagans and Lasiurus cine- reus.
On 27 July 1967, I captured a female that carried two embryos (8 in crown-
rump length) in the attic of a classroom building on the campus of the Black Hills State College at Spearfish. Females ordinarily give birth to one or two young between May and mid- June in western South Dakota (e.g., a female taken in Fall River County on 14 June carried two embryos that each measured 7 in crown-i-ump length ) ; thus a pregnant big brown bat in late July would seem ex- ceptional. Returning to the same attic on 29 July I obtained 21 additional E. fusciis, of which eight were lactating adult females, five were subadult fe- males, seven were subadult males, and one was an adult male.
Martin (pers. com.) reported that a maternity colony of this species was lo- cated on 19 May 1970 in the attic of the Dakota Steel Company in Rapid City. Two other maternity colonies were found in Hot Springs in the summer of 1968. Ten pregnant females (six were banded and released, and four were ex- amined; each of the latter carried a single embryo of an average crown-rump length of 18.5 (15.0-23.0)) and four lac- tating females were netted in the attic of the Fall River Museum on 25 June and 27 July, respectively. One subadult male and four subadult females were captured in the Fall River County Court- house on 26 July.
A captive female from the Fall River Museum aborted two young on the night of 25 June (Turner and Davis, 1970: 363 ) . The entire process lasted less than 10 minutes. The female propped herself on the tips of her forearms, lowered her head ventrally toward the vaginal ori- fice, and actively aided the birth by licking the vaginal region and the emerg- ing young. The young were cradled in the uropatagium, which was curved for- ward beneath the body of the female. Though seemingly close to exhaustion, the female readily devoured several moths offered to her immediately follow- ing the events just described.
Length of testes of 18 adult males ob- tained between 13 June and 4 July were 5.8 (4.0-7.5). Testicular length of two
TURNER: MAMMALS OF THE BLACK HILLS
55
adults taken on 29 Jul)' 1967 were 9 and 10, whereas that of five subadults cap- tured on the same day were 4.4 (3.0-7.0). A male (banded and released), with large, descended testes, was extracted from a space between a stone chimney and eaves of an overhanging roof of a house in Rapid City on 3 August 1969. Jones and Genoways (1967b: 191) described molt in E. f. juscus and their description seems applicable also to E. f. pallidiis. Subadults from Spearfish and Hot Springs were undergoing post- juvenal molt late in July, whereas an adult female from Spearfish was in proc- ess of annual molt.
Big brown bats from southwest of Minnekahta harbored bat bugs, Cimex adjunctus Barber, and ticks, Ornitho- doros kelleyi Cooley and Kohls, whereas a female from Hot Springs was infested ' with spinturnicid mites, Spinturnix hakeri Rudnick, and a male from Moon shel- tered bat flies, Basila forcipata Ferris. Eptesicus from various localities in the Black Hills were parasitized externally by chiggers, Leptotrornbidhitn imjotis (Ewing).
Specimens examined (129).— SOUTH DA- KOTA: Lawrence County: Spearfish, 3600 ft, 22; Little Spearfish Canyon, Savoy, 8 mi W Lead, 5200 ft, 2; 2 mi W Nemo, 4700 ft, 6; 1 mi E Nemo, 4700 ft, 1. Pennington County: Rapid City, 5 (4 NRW, 1 SDMT); Diamond S Ranch, near Rapid City, 6 (UMMZ); South Canyon, 3 mi W Rapid City, 12 (AMNH); Moon, 22 mi W Hill City, 6200 ft, 1; Ice Cave, 8 mi N, 15 mi W Custer, 6400 ft, 1; Hill City, 1 (NRW); 3 mi SE Hill City, 5300-5400 ft, 3 (UMMZ); 10.5 mi E Wyoming border, 5.8 mi N Custer County line, 2 (MSWB). Custer County: 5.75 mi N, 5.75 mi E Custer, 5220 ft, 5; Hell's Canyon, 13 mi W Custer, 2 (USNM); Custer, 4 (USNM); Squaw [Grace Coolidge] Creek, 1 (AMNH); French Creek Cave, Custer State Park, 8 (SDMT); Jewel Ca^■e, 2.5 mi S, 12 mi W Custer, 5280 ft, 2 (1 UW, 1 WCNP); Headquarters, Wind Cave National Park, 4100 ft, 17 (2 WCNP, 15 UMMZ); Shirttail Canyon, Wind Cave Na- tional Park, 2 (UMMZ); Ony.x Cave, 10 mi NW Hot Springs, 1 (UW). Fall River County: Fall River Museum, Hot Springs, 3600 ft, 8; Courtliouse, Hot Spnngs, 3500 ft, 5; 5.5 mi E Minnekahta, 4000 ft, 1; 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 8.
WYOMING: Crook County: Sand Creek,
1 (USNM). Weston County: 1.5 mi E Buck- horn, 6150 ft, 1; Cambria Coal Mine, 6 mi N Newcastle, 6000 ft, 1.
Additional record.— SOUTH DAKOTA: Meade County: Fort Meade (Miller, 1897:98).
Lasiurus borealis borealis (Miiller) Red Bat
Vespertiliu borealis Miiller, 1776, Des Ritters Carl von Linne . . . vollstandiges Natm- system . . ., suppl., p. 20 (type locality. New York ) .
Lasiurus borealis — Miller, 1897, N. Amer, Fauna, 13:105, 16 October.
Turner and Davis (1970:363) re- cently reported the first specimens of the red bat to be taken west of the Missouri River in South Dakota. Jones and Geno- ways (1967b: 193) suggested that this species probably reached the western limits of its resident range in South Da- kota at approximately the 101st meridian, but that migrating individuals might pass through the western part of the state on their way to or from regions farther to the north. Long (1965:534) mentioned an individual (USNM 5264) that was hsted by H. Allen (1864:20, 1894:153) from Laramie Peak, as the only specimen known from Wyoming.
Nets were established over two ponds near Moon Campground in Pennington County on 29 July 1968. One net was located in a small meadow in a woodland and the other over a farm pond situated in a pasture. No red bats were taken over the woodland pond, but Wayne H. Davis captured a female young of the year over the farm pond; the latter indi- vidual was just beginning post-juvenal molt. Two Mijotis leibii, two M. keenii, two M. volons, two Lasiurus cinereus, and two Lasionycteris noctivagans were removed from this same net, whereas four M. keenii, one M. lucifuf^us, two M. fJiysanodes, three M. volans, four E. fus- cus and 13 L. cinereus were removed from the net over the woodland pond. Returning to the woodland pond on 20 August, I obtained a lactating female red bat, along with one M. keenii, one M. Jucifufi^us, one M. thysonodes, two M. volans and one L. cinereus. On the first
56
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
visit, the woodland pond was fairly open and activity began about 21 hrs (MDT); a total of 27 bats was netted. In contrast, the pond was well overgrown with vege- tation on the second visit. Although the net was observed from 20 until 00:30 hrs, not a single bat was detected; how- ever, when examined early the next morning, the seven bats in the net were concentrated over a small pool of open water. Utilization of the pond by bats evidently decreased greatly as the sur- face vegetation increased.
The interim (22 days) between cap- ture of the flying young of the year and the lactating female almost certainly ex- cludes the possibility that the young was that of the female. Thus, at least two families of L. borealis evidently occurred in the vicinity of Moon during the sum- mer of 1968. Members of this species migrate southward in winter, returning northward again in spring, and addi- tional data are needed to ascertain the summer status of the red bat in the Black Hills.
Specimens examined (2). — SOUTH DA- KOTA: Pennin0on County: Moon, 22 mi W Hill City, 6200 ft, 2.
Lasiurus cinereus cinereus
(Palisot de Beauvois)
Hoary Bat
Vespertilio cinereus (misspelled lincreus) Pali- sot de Beauvois, 1796, Catalogue raisonne du museum de Mr. C. W. Peale, Phila- delphia, p. 18 (p. 5 of English edition liy Peale and Palisot de Beauvois ) ( type lo- cality, Philadelphia, Pliiladelphia Co., Penn- sylvania).
Lasiurus cinereus — H. Allen, 1864, Smiths. Misc. Coll., 7 ( 165 ):21, June.
The hoary bat has not been reported from the Wyoming section of the Black Hills, but it is known from Lawrence, Pennington, Custer, and Fall River coun- ties of South Dakota. This is the largest of the Black Hills bats and seems to be locally abundant in suitable habitats. Like the red bat, Lashinis cinereus is a migrant, arriving in the Hills region in late spring and residing there until early
autumn; seasonal records are from 13 June to 31 August.
Findley and Jones (1964:469) indi- cated that females generally precede males in the northward migration and occupy the plains and adjacent lowlands in warmer months, whereas the males usually are found at higher altitudes or, occasionally, at higher latitudes. The 28 specimens (11 males and 17 females) taken in this study certainly establish the Black Hills as one of the few areas in the mid-continent region where adult L. cinereus of both sexes occur together in summer. In western South Dakota, males occur only in the Black Hills, at eleva- tions of 4700 to 6200 feet, whereas fe- males occupy suitable habitat up to 6400 feet in the Hills and also range down through transitional lower elevations out onto the adjacent plains and lowlands. The first representatives of this species from the Black Hills were reported by Jones and Genoways ( 1967b : 193 ) .
Of 14 hoarv bats collected in the vicinity of Nemo between 28 June and 3 July 1967, seven were netted over Box- elder Creek (see account of M. Jucifu- gus ) , and seven others were shot in areas where M. leibii, M. hicifugus, M. voJans, E. fusctis, and Lasionycteris noctivagans also foraged (i.e., over streams and mea- dows bordered by ponderosa pine). A single male was shot on 19 June 1967 as it foraged over a small valley northeast of Custer, and two females were obtained (one netted over a stock pond, in asso- ciation with the above mentioned species and M. Hu/sanodes, and the other shot as it flew along the edge of a conifer- bordered meadow) southwest of Minne- kahta on 13 June 1967. Of 15 hoary bats ( 1 1 banded and released ) netted over a woodland pond (13 individuals) and a farm pond (two individuals) near Moon on 30 July 1968, six were adult males, six were lactating females, and three were juveniles (one male and two females); an additional male young of the year was obtained o\'er the wood- land pond on 20 August 1968 (see ac- count of L. borealis). Two female L.
TURNEH: MAMMALS OF THE BLACK HILLS
57
cinereus (one lactating and the other a young of the year) were netted over a sewage setthng pond at the head of Wind Cave Canyon on 25 July 196(S in company with a L. noctivag,ans.
One or (usually) two young are born from late May to early July. Of two adult females taken on 13 June, one car- ried two embryos (each being 22 in crown-rump length) and the other was lactating. None of six females taken be- tween 28 June and 3 July was pregnant, but four were lactating; all six adult females obtained at Moon on 29 July, and one collected in Ditch Creek Valley on 13 June, also were lactating. Testes of eight males captured from mid-June to mid-July were 5.4 (4-7) in length.
The subadults collected late in July to mid-August were in fresh pelage. No ectoparasites were found on individuals of this species.
Specimens examined (28). — SOLTTH DA- KOTA: Lawrence County: 2 mi E Tinton, 5400 ft, 1; 2 mi W Nemo, 4700 ft, L3; 1 mi E Nemo, 4700 ft, L Pennington County: Rapid Citv, 1 (NRW); Moon, 22 mi W Hill Cit>-, 6200 ft, 5 (2 UK); Ditch Creek, 14 mi W Hill City, 6400 ft, 1; 10.5 mi E Wyoming border, 5.8 mi N Pennington-Custer county line, 1 (MSWB). Custer Counti/: 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; Wind Cave Can- yon, Wind Cave National Park, 4100 ft, 2. Fall River Countti: 0.5 mi S, 1.5 mi W Minne- kahta, 4200 ft, 2.
Piece tus townsendii palleseens (Miller) Townsend's Big-eared Bat
Corynorhinus macrotis palleseens Miller, 1897, N. Amer. Fauna, 13:52, 16 October (type locality, Keam Canyon, Navajo County, Ari- zona).
Plecotus townsendii palleseens — Handley, 1959, Proc. U. S. Nat. Mus., 110:190, 3 Septem- ber.
This cavernicolous species is distrib- uted through much of western North America, and is known from all counties that comprise the Black Hills; it was first reported from the study area by G. M. Allen (1916:334) under the name Cory- norhinus megalotis palleseens. John Ty- ers and associates of the U. S. National Park Service have banded many individ- uals in Jewel Cave, where an estimated
2000 big-eared bats hibernate in winter along with Mijotis leihii, M. liieifugus, M. volans, and Eptesieus fiiscus. Bats evidently vacate these limestone hiber- nacula in late spring and disperse throughout the Hills; females congregate in maternity colonies in warm standstone caves or attics of buildings, whereas males seem to remain solitary. Known altitudinal range in the Hills is from 3400 to 6200 feet.
Although large associations of big- eared bats occur in the Hills region, none of the specimens taken in this study was obtained by the usual collecting methods, that is, by netting or shooting. Perhaps the activity patterns of this bat differ from those of other species that occur in the region, and it may remain in re- treats until well after dark. V. Bailey, however, reported P. townsendii to be among the earliest observed bats flying each evening in the vicinity of Sundance, Wyoming (Handley, 1959:175).
On 4 August 1969, a solitary big-eared bat was observed in a narrow chimney of a small cave on the rim of Little Spearfish Canyon, above Timon Camp- ground; a single male (testes 6 in length) was captured in the cave on 7 August 1970. Although the cavern is used exten- sively as a night roost (see account of M. leihii), P. t. palleseens were the only bats that utilized the chimney as a day roost.
On 20 July 1967, I located a nursery colony in a small cave formed in a sand- stone bluff at a locality 1 mi S and 2 mi E Hot Springs, Fall River County (Turner and Jones, 1968:447). Of the 41 bats present in the cavern, 22 were adult females; 19 of these were lactating and one carried an embryo that was 27 in crown-Runp length. Eight juvenile fe- males and 11 juvenile males (with tes- ticular lengths of 3) comprised the re- mainder of the colony. The young rep- resented various stages of development and molt (from fine post-natal hair on nearly naked animals to juvenal pelage on well de\'eloped }'oung) indicating that dates of parturition within the col-
58
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
ony probably extend over a considerable period of time. A solitary male (testes 6 in length) was found in this same cave on 30 June 1968.
A secluded male (testicular length 4) was found clinging to the crystalline cal- cite (dog-tooth spar) walls of Daven- port Cave on 20 November 1967, and three big-eared bats ( two males and one female) were found hibernating in an unnamed limestone cave in Dark Can- yon on 10 November 1967 (see account of L. noctivagans) . Although the males were captured deep in the cave, the solitary female was taken in the twilight zone. Testicular length of both males was 3. Ambient temperatures at the sites of capture were 6.4°, 5.3°, and 7.2 °C, whereas the rectal temperatures of the bats were 7.2°, 8.0°, and 8.6°C, respec- tively. Mummified remains of P. toicn- sendu were found in Jewel Cave on 13 June 1965, and in the deserted Cambria Coal Mine complex north of Newcastle, Weston County, Wyoming, on 2 July 1965.
Jewel Cave was visited by a field party from the Museum of Natural His- tory on 20 November 1967. Approxi- mately 600 Plecotus were wintering there at the time, and 304 of these were examined. Two hundred big-eared bats (69 females and 131 males) were distrib- uted in clusters containing from two to 33 bats; the remaining 104 individuals ( 40 females and 64 males ) were solitary. Ambient temperature ranged from 5.0° to 6.4°C and the relative humidity ranged from 64 to 70 percent in areas occupied by Plecotus. The average rec- tal temperatures of 18 females and 34 males were 4.3°C (3.6-6.2) and 4.2°C (3.6-6.2), respectively; individuals from larger clusters tended to have a mean body temperature 1-2° lower than did solitary individuals or those from small clusters. Most of the bats were within 150 feet of the entrance, where the am- bient temperature was lower than deeper in the cave. Air currents made this 2:en- eral area susceptible to environmental fluctuations.
A plug that completely filled the
vaginal orifice was observed in many of these females when rectal temperatures were taken. Thomas H. Kunz examined seven vaginal smears under a phase con- trast microscope and found numerous agglutinated clumps of sperm in the semen suspension. Pearson et ah ( 1952: 293) indicated that vaginal plugs were not found in Plecotus townsendii studied in California.
Jones and Genoways (1967b: 194) noted the complete absence of big-eared bats in Jewel Cave on 12 June 1965, and speculated that Plecotus vacates the cave entirely in late spring and for most of the summer. However, nets placed within the entrance to Jewel Cave on 24 July 1968 captured four Plecotus along with several other species (see account of M. hicifugus). Of the 2165 big-eared bats banded by the National Park Service at Jewel Cave from December 1959 through December 1963, nine returns have been reported. Four of these were taken within an area 30 miles south and east of Jewel Cave between elevations of 3400 and 3800 feet (see additional records) during the warm months (5 May to 1 September ) . The others were recaptured within the cave itself in winter. A male that was banded on 31 December 1959 was recaptured in Jewel Cave on 24 July 1968; thus, there was an interim of 3148 davs between the time of banding and recapture (Turner and Davis, 1970: 364).
Of those specimens examined in Jewel Cave on 20 November 1967, 67 percent were parasitized by bat flies, Trichohius corynorliini Cockerell and one individual harbored a chigger, Lep- totromhidium mijotis (Ewing). Sarcop- tid mites, Sarcoptes lasionycteris Boyd and Bernstein, were prevalent also in the Jewel Cave population. Female big- eared bats displayed a higher incidence of parasitism than did males. Two fe- males from the maternity colony near Hot Springs, were parasitized by the same species of bat fly and by laelaptid mites, Macronyssus longisetosus ( Fur- man ) , whereas a female from Dark Can-
TURNER: MAMMALS OF THE BLACK HILLS
59
yon harbored a different species of lae- laptid mite, Macronyssus iinidens Ra- dovsky, representing tlie first record of this parasite in Soiitli Dakota.
One of tlie 22 adult females from the nursery colon) near Hot Springs (ob- tained on 20 July 1967) was molting simultaneously over both dorsum and venter.
Specimens examined (168).— SOUTH DA- KOTA: Lawrence Coiuitt/: Cleopatra Mine, 1.5 mi N Carbonate, 2 (SDMT); T. 5 N, R. 2 E, NE }, sec. 17, 2 (SDMT); "near Chicken Ranch and Rouliaix Lake," 1 (NRW); Crow- feet [=Cro\v Peak?], Black Hills, 7 (AMNH); R. B. Hayes Mine, 1 (SDMT). Meade County: Da\enport Cave, 3 mi S, 0.5 mi W Sturgis, 4400 ft, 1. Pennington County: Tool Shed Cave, N Battle Creek, S Haywood, 3570 ft, 8; Rochford, 5300 ft, 1 (UMMZ); Deadman Can- yon, 2-3 mi W Rapid City, 1 (NRW); Rapid City, 1 (NRW); Dark Canyon, 2 mi S, 5 mi W Rapid City, 3800 ft, 3; Spring Creek Canyon, 1 (NRW); 4 mi SW Rockerville, 5000 ft, 1 (UMMZ); Igloo Cave, 3 mi S, 16 mi W Hill Cit>-, 12 (SDMT); 3 mi SE Hill City, 3 (UMMZ); Bear Trap Cave, 4 mi S, 16 mi W Hill City, 6200 ft, 2 ( 1 SDMT). Custer County: Hell's Canyon, 13 mi W Custer, 7 (USNM); Cuvahoga Mine, 2 (SDMT); 18 mi W Custer, 1 (USNM); Jewel Cave, 2.5 mi S, 12 mi W Custer, 5280 ft, 52 (2 NRW, 3 SDMT, 1 UW, 4 WCNP); S and G Cave, T. 5 S, R. 3 E, NW }i, NW }i sec. 3, 2 (SDMT); 2 mi W Wind Cave National Park, 1 (UW); Wind Cave, Wind Cave National Park, 2 (1 SDMT, 1 UW). Fall River County: 1 mi S, 2 mi E Hot Springs, 3400 ft, 41; 5.5 mi E Minnekahta, 4000 ft, 1.
WTOMINC: Crook County: Sand Creek, 10 mi E Sundance, 3750 ft, 10 (USNM). Weston County: 4 mi E Four Corners, 1; 4 mi N, 3 mi W Newcastle, 1.
Additional records.— SOUTH DAKOTA: Custer County: Custer (G. M. Allen, 1916: 344); French Creek Cave, Custer State Park (R. A. Martin, pers. com.); 4 mi W Fairhurn (band record: Tyers, 1963:17). Fall River County: 2 mi N, 1 mi W Burdock (band rec- ord); Cascade (band record); Cascade Springs ( band record ) .
ORDER LAGOMORPHA— Hares, Rabbits, and Pikas
Family LEPORIDAE— Hares AND Rabbits
Four species of lagomorphs, one of the genus Lepus and three of the genus Sylvila<j,us, are native to the Black Hills; two additional species of the genus
Leptis may occur in the immediate vi- cinity of the Hills. The four native leporids represent interesting examples of similar species with either parapatric or sympatric, but ecologically distinct, distributions in the Hills region.
Sylvilagus audubonii baileyi (Merriam)
Desert Cottontail
Lej)us baileyi Merriam, 1897, Proc. Biol. Soc. Washington, 11:148, 9 June (type locality, Spring Creek, E side Bighorn Basin, Big Horn Co., Wyoming).
Sylvilagus auduhonii haileyi — Lantz, 1908, Trans. Kansas Acad. Sci., 22:236.
Nelson (1909:234) first reported the desert cottontail from the Black Hills on the basis of a young female obtained at Elk Mountain by Merritt Gary on 10 July 1906. Sij]vi]a(i,us auduhonii is a common mammal on the western Great Plains and inhabits grasslands up to about 5000 feet in the Hills. Specimens are available only from Pennington, Cus- ter, and Fall River counties, although the species undoubtedly occurs along the entire interface of the Black Hills and surrounding prairie. The desert cotton- tail is geographically sympatric with the eastern cottontail and with Nuttall's cot- tontail; however, these three rabbits usu- ally occupy different ecological niches.
In the western section of the Hills, Nuttall's cottontail has been collected in the same general area with the desert cottontail. Where the two occur to- gether, S. nuffaUii usually occurs at higher elevations in wooded or brushy habitat, whereas S. auduhonii lives in relatively open, grassy situations at lower elevations. For example, Merritt Gary obtained both species at Elk Mountain early in the 1900's; S. nuttaUii was taken at 6000 feet, and S. auduhonii at 5000 feet. A similar ecological separation ap- parently exists between S. auduhonii and S. jioridanus in the eastern section of the Black Hills (see account of S. jioridanus) .
Four females taken near Minnekahta in mid- June 1967 were shot in arid grass- land and pastures. Bluegrass, brome grass, cord grass (Spartina pectinata), western wheat grass, sagebrush, soap-
60
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
weed, pricklypear cactus, and various mustards {Brassica sp.) comprised the lowland vegetation. Syhilagus audu- honii is common in Wind Cave National Park where upland prairie is the pre- ferred habitat. In addition to the above- mentioned plants, big bluestem, little bluestem, blue grama, sideoats grama, green needlegrass, needle and thread grass, buffalo grass, yellow sweet clover, blue vervain, beardstongue (Penstemon alhidus), wavyleaf thistle (Cirsium tin- dtilatiim), bluestem pricklypoppy (Ar- g,emone poh/anthemos), western wall- flower {Erysimum asperum), blackeyed susan {Rudheckia hirta), yellow cone- flower {Ratihida columnifera), puiplc coneflower (Echinacea anfi,ustifolia) , goosefoot {Chenopodium sp. ), common sunflower {Helianthus anniius), scarlet globemallow (Sphaeralcea coccinea), wild alfalfa, woolly mullein (Verhascum thapsus), common milkweed (Asclepias syriaca), and mariposa lily clothe the prairie portion of the Park inhabited by desert cottontails.
Cranially and externally, S. auduhonii frequently is difficult to differentiate from S. rmttallii. Both species have large audi- torv bullae and a large external auditorv meatus, but in S. auduhonii the mesop- terygoid fossa and supraoccipital shield are slightly differently shaped and the pelage is somewhat paler than in S. nuttallii. A female cottontail from 3 mi E Custer reported by Stebler ( 1939:390) as Sylvi1a<s,us nuttallii p^rana^eri is best assigned to Sylvihip,us audid)onii hailcyi. This particular specimen is intermediate in all cranial characters, but has the paler pelage of the desert cottontail. The individual represents the greatest pene- tration of this species into the wooded interior of the Black Hflls; however, it was taken in an area that ". . . appears to be a natural prairie, for no evidence indicating the presence of a former for- est was discovered" (loc. cit.). Sage- bmsh, goldenrods, butter and eggs ( Lin- eria vulgaris), and asters were conspicu- ous among various grasses at the site of capture.
Of four females shot southwest of Minnekahta in mid-June, one carried six embryos, another was lactating; the two others were juveniles in fresh, fine juvenal pelage. Subadult desert cottontails are present in collections made from June to September. A female taken near Custer on 16 August 1934 was in fresh summer pelage, whereas another obtained in \\^ind Cave National Park on 7 August 194S was molting simultaneously over both dorsum and sides. An adult female captured at the latter localit)' on 1 April 1950 still was in well-worn winter pelage. An adult female and a juvenile female obtained near Minnekahta both were parasitized externally by ticks, Haema- physalis leporispalustris (Packard).
Specimens examined (18). — SOUTH DA- KOTA: Pennington Counhj: Rapid Citv, 2 (NRW); Rockerville, 1 (UMMZ). Custer County: 4 mi E Custer, 1 (UMMZ); BufFalo Corral, Wind Cave National Park, 4300 ft, 1; Curh- Canyon, 3 mi N Wind Cave, Wind Cave National Park, 1 (WCNP); 2 mi S Wind Cave, Wind Cave National Park, 2 (WCNP); Shirt- tail Canvon, Wind Ca\e National Park, 4 (UMMZ); Elk Mountain, 5000 ft, 1 (USNM). Fall River County: 0.5 mi S, 1.5 mi W Minne- kahta, 4200 ft, 4; 1 mi N, 4 mi E Edgemont, 1.
Sylvilagus floridanus similis Nelson
Easterx Cottontail
Sylvilagus floridanus similis Nelson, 1907, Proc. Biol. Soc. Washington, 20:82, 22 July (type localit>', Valentine, Cherry Co., Ne- braska ) .
Sylvilagus floridanus occupies ripar- ian communities and brushy draws across tlic Great Plains and into the foothill transition zone of the Black Hills. East- ern cottontails are uncommon in the Hills; the>^ occur below 4200 feet in the eastern section, where they are sjmpatric with, but ecologically distinct from, S. auduhonii. Sylvilagus floridanus has been reported previously from the study area as Lepus sylvaticus by Bailey (1888: 446), who noted that this rabbit is com- mon in bushy ravines in the northeastern part of the Hills; however, specimens are available only from Lawrence, Custer and Fall Ri\'cr counties.
\\^hile a Ranger-Naturalist at Wind
TURNER: MAMMALS OF THE BLACK HILLS
61
Cave National Park in tlic suninicr of 1968, I had occasion to witness contrast- ing habitat selection between S. /. similis and S. a. Inulciji. Eastern cottontails are confined to brnshy draws, forest edges of \'alley systems that infringe into the foot- liills, and thickets along sides of streams; desert cottontails frequent the open prairie and upland grasslands. Several mornings were spent observing both spe- cies feeding within five feet of one another on a closely mowed lawn near Park Headquarters. To the north of the lawn was a brushy ravine of mountain mahogany, wild rose, hackberry {Celtis occidentalis), and chokecherry; all else was upland prairie. When startled, S. fioridanus sought refuge in the ravine, whereas S. auduhonii behaved much as a jackrabbit and loped off across the grassland. At the north end of the ravine is Wind Cave proper; I found the skele- ton of an eastern cottontail therein on 23 August 1967. Apparently, the rabbit had been washed into the cave subse- quent to a heavy rain because bones were found about 80 feet below the surface, where there was no vegetation or signs of active inhabitation by this species.
A female taken in the Park on 22 July showed no signs of reproductive activity, and a male obtained there on 31 August was a juvenile that was parasitized ex- ternally by ticks, Ixodes spinipaJpis Had- wen and Nuttall, and chiggers, Eutrom- bicula alfreddugesi (Oudemans). Two adults, a male and a female, taken near Hot Springs in April 1950 by J. A. King, were molting over the rump.
Specimens examined (6). — SOUTH DA- KOTA: Lawrence County: 7 mi N Spearfish, 1. Custer County: Wind Cave, Wind Cave National Park, 1; Headquarters, Wind Cave National Park, 4100 ft, 2. Fall River County: 2 mi N Hot Springs, 1 (UMMZ); 1 mi S Hot Springs, 1 (UMMZ).
Additional record.— SOUTH. DAKOTA: Pennington County: Rapid City (Bailey, 1888: 446).
Sylvilagus nuttallii grangeri (J. A. Allen)
Nuttall's Cottontael
Lepus sylvaticus grangeri J. A. Allen, 1893, Bull. Amer. Mus. Nat. Hist., 7:264, 21 Au-
gust (type locality. Black Hills, Hill City, Pennington Co., South Dakota).
SylvUagus nuttallii grangeri — Nelson, 1909, N. Amer. Fauna, 29:204, 31 August.
Sylvilagus nuttallii grangeri, orig- inally described from the Black Hills, occupies areas of sagebrush and timber in the montane western United States. In the Hills, this species occurs in spruce and pine forests of the boreal cap, down through the transitional foothill zone to about 4500 feet; at the lower elevations in the western section of the Hills, forest edges and brushy draws are inhabited. In such ecotonal areas, S. nuttallii is sym- patric with S. audidjonii, from which it often is difficult to distinguish (see ac- count of this species). Albeit the most abundant lagomorph in the Black Hills, specimens of Nuttall's cottontail are lack- ing from Meade, Fall River, and Weston counties.
I know of no instances of parapatry between S. nuttallii and S. fioridanus in the study area, although such has been reported from Laramie County, Wyoming (Hall and Kelson, 1951:53). Based on the specimens examined, the nearest lo- calities of record between these two lago- morphs in the Black Hills proper are approximately 15 miles distant [S. fiori- danus from Wind Cave National Park (4100 feet), and S. nuttallii from Custer (5300 feet)]. Whereas Nuttall's cotton- tail inhabits wooded environs above 4500 feet in the central and western Hills, the eastern cottontail occupies brushy ra- vines and streamside thickets below 4200 feet in the eastern Hills and along the northern and southern peripheries. Both species are distributed outward from the Hills along riparian communities on the plains, S. fioridanus to the east and S. nuttallii to the west. Intergradation between nuttallii and fioridanus has been suggested as possibly occurring along the eastern base of the Rocky Mountains, but Hall and Kelson (loo. cit.) concluded that this is not the case, partially on the basis of a study of specimens from the Black HUls.
Individuals from southeast of Hill
62
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
City, Pennington County, were collected from pine-clad uplands and spnice-filled canyons, usually near an abundance of rocks, fallen logs, shrubbery, or old buildings, all of which are in close prox- imity to small open grassy areas. On 7 July 1967, a melanistic juvenile which had sought refuge under a ponderosa pine log was captured on a wooded hill- side near Nemo, Lawrence County.
A female obtained in Palmer Gulch, Pennington County, on 12 April 1946 carried three embryos that were 29 in crown-rump length; another taken near the same site on 13 August 1950 con- tained four embryos that were 17. A lac- tating female was captured on 10 Au- gust 1929 at Elk Mountain, and a preg- nant female containing five embryos that were 56 ( crowni-rump ) was shot north- east of Custer on 20 June 1967. Juveniles and subadults of this species are common from mid-April to early September.
The holotype, obtained on 10 August 1894 by W. W. Granger, is a nearly full- grown young of the year in subadult ( post-juvenal ) pelage; whereas, four paratypes are juveniles in fresh juvenal pelage, and another is an adult with worn pelage that was commencing to molt. New hairs can be observed under- lying the older pelage on the rimip of a topotype obtained on 25 April 1910, and on adults generally taken in June and early July. Apparently, adults molt to summer pelage from late March to mid- July, whereas change to winter pelage begins early in August on some individ- uals. The melanistic juvenile is entirely black except for diffuse ochraceous guard hairs over the sides, venter, and white hairs along edges of the ears and on the tail.
Specimens examined (50). — SOUTH DA- KOTA: Lawrence County: 2 mi S Tinton, 6100 ft, 1; 2 mi W Nemo, 4700 ft, L Penning- ton County: Hill City, 7 (1 USNM, 6 AMNH); Redfem, 10 (USNM); Palmer Gulch, 3-4 mi SE Hill City, 5300-5400 ft, 15 (9 UMMZ); 16 mi NW Custer, 2 (UMMZ); near Mount Rushmore, 1 (UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, 1 (FMNH); 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; Custer, 3 (USNM); Bull Springs, 2 mi N, 9 mi W
Custer, 1 (UMMZ); Elk Mountain, 2 (1 USNM, 1 UMMZ); Redbird Canyon, Elk Mountain, 6000 ft, 1 (USNM).
WYOMING: Crook County: De\ils Tower, 3 (USNM); Sundance, 1 (USNM).
Lepus townsendii campanius Hollister
\\^HITE-TAn.ED JaCK RaBBIT
Lcpus campestris Bachman, 1837, J. Acad. Nat. Sci. Philadelphia, 7:349 (type locality, plains of Saskatchewan, probably near Carl- ton House ) .
Lepus townsendii campanius — Hollister, 1915, Proc. Biol. Soc. Washington, 28:70, 12 March, a renaming of Lepus campestris Bachman.
^^ hite-tailed jack rabbits inhabit up- land grasslands of the northern plains and montane regions of western United States. Abundant on the surrounding flats, Lepus townsendii is less common in the Black Hills proper. It inhabits open woodlands up to 6000 feet and iso- lated grassy areas such as Reynolds Prairie, Gillette Prairie, and the Bald Hills. Although no specimens are avail- able from Custer and Fall River counties, this species often has been observed along roads at night in these areas. Bailey (1888:438) recorded the first rep- resentatives of L. townsendii from the Hills (as L. campestris), and Palmer (1897:28, 74) indicated that a single hunter near Newcastle, Weston County, killed more than 100 indi\iduals in 1893-94.
At lower elevations, these lagomorphs inhabit open prairie and upland grass- land. I have observed them in Wind Cave National Park and in pastures just southeast of Minnekahta; in such situa- tions L. tonnsendii o\erlaps ecologically with SyIvilaL!,tis aiidu1)onii. At higher elevations, L. townsendii frequents open woodlands of aspen, pine, and spruce that border meadows and pasturt\s; un- der these conditions, L. townsendii over- laps ecologically with Sylvikig,tis nuttaUii to an undetermined extent.
A female shot southeast of Hill Cit\' on 12 April 1946 contained four embryos that were 70 in crown-rump length; two males taken on 15 June 1965 and 28 June 1967 had testes that were
TURNER: MAMMALS OF THE BLACK IIILLS
63
49 and 52 in length, respectively. Yonng of the year were present from mid-March to late August; in 1894 Palmer (1897:28) observed parturition of individuals at Newcastle, Wyoming, on 23 and 24 May. Specimens obtained throughout the colder months were in white pelage [V. H. Cahalane ( field notes, on file at Wind Cave National Park, p. 66) observed an indi\idual in which the change to the white winter coat was well-ad\'anced on 3 October 1934]. The presence of new brownish hairs beneath the well-worn white hairs indicated that the pregnant female mentioned above was undergo- ing molt from winter to summer pelage. A male shot southwest of Sturgis was infested with ticks, Dermacentor ander- soni Stiles.
Specimens examined (17). — SOUTH DA- KOTA: Lawrence County: Deadwood, 1 (US\M). Meade County: Fort Meade, 1 (USNM); Black Hawk, 1 (NRW); Vanocker Canyon, 7 mi S, 1 mi W Stnrgis, 4800 ft, 1. Pennington County: Rapid Cit>', 3 (USNM); 3.5 mi i\, 1.5 mi E Deerfield, 1; Hill City, 1 (UMMZ); 3 mi SE Hill City, 5300 ft, 2 (UMMZ); 3 mi W Rockerville, 1 (UMMZ).
WYOMING: Crook County: Devils Tower, 1 (USNM); Sundance, 6000 ft, 1 (USNM). Weston County: Newcastle, 3 (USNM).
ORDER RODENTIA— Rodents
Rodents are by far the most numerous of Black Hills mammals, both in number of kinds and in number of individuals. Seven families of rodents, totaling 22 species of 18 genera are indigenous to the study area; two families ( represented by three genera, each of which contains but one species) have been introduced into the Hills. Four species of rodents (representing two additional genera) ha\'e been incorrectly reported from the region, and five species (representing two additional genera) may possibly occur in the Black Hills. On the basis of the numbers of individuals and resultant biomass, rodents are the base of the vertebrate food-chain in the Hills.
Family SCIURIDAE— Squirrels AND Allies
Sciurids in the Black Hills represent
a di\ersitv of lilc lorms and habits. Among terrestrial squirrels, three are fos- sorial (Cijnomys, Spermophilus, and Mannota) and one is scansorial {Eutarn- ias); two arboreal squirrels are scansorial (Sciurus and Taniiasciurus) and one is volant (Glauconiys). Except for noc- turnal flying squirrels, all other taxa are diurnal and are among the most con- spicuous members of the Black Hills mammalian fauna. Seasonal activity varies for the different species, some hi- bernate in colder months, whereas others remain active the year around.
Eutamias minimus pallidas (J. A. Allen)
Least Chipmunk
[Tamias quadrivitatus] var. palUdus J. A. Allen, 1874, Proc. Boston Soc. Nat. Hist., 16:289, June (type locality restricted to Camp Thorne, near Glendive, Dawson Co., Mon- tana, by Cary, Proc. Biol. Soc. Washington, 19:88, 4 June 1906).
Eutamias lyiinimus pallidus — A. H. Howell, 1922, Jour. Mamm., 3:183, 4 August.
This pale chipmunk is widely distrib- uted in the western section of the North- ern Great Plains, and inhabits riparian and scarp woodlands as well as shnab- land habitats. In the Black Hills, E. m. pallidus occurs in the forested or brushy ravines of the lower southeastern foot- hills. Red Valley, and adjacent plains, at elevations up to 4300 feet. Specimens from Wind Cave National Park and from southwest of Minnekahta clearly are re- ferrable to pallidus, whereas all other chipmunks examined from the Black Hills proper are assignable to the en- demic race, silvaticus (see the following account). Distribution of E. minimus represents one of two examples of two subspecies of the same mammalian spe- cies occurring within the defined bound- aries of the Black Hills (see account of Dipodomys ordii). Sutton and Nadler (1969:526) described two karyotypes (A and B) in the genus Eutamias, and indi- cated that absence of heteromorphic chromosomes is evidence that the sub- species characterized by these two dif- ferent karyotypes do not intergrade. Eu-
64
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
tamias minimus pciUidus and E. m. sil- vaticus both possess karyotype B; speci- mens of intermediate characteristics are encountered in the lower foothill zone, near Elk Mountain and Sundance for example.
Little ecological data are available for E. m. pallidus in the Black Hills re- gion. Individuals are extremely abun- dant along the slopes of Wind Cave Canyon, which is vegetated by mountain mahogany, wild rose, hackberry, choke- cherry, wild plum, pricklypear cactus, pricklypoppy, and soapweed. V. H. Ca- halane observed this small sciurid con- suming dried wild cherries and wild plums (field notes on file at Wind Cave National Park). On 29 July 1968, a fe- male olive-backed pocket mouse cap- tured in a snap-trap was partially devoured by a chipmunk that departed as I approached. Perognathus jasciatus, P. Jiispidiis, Pewmyscus manicidatus, Reithwdontomys megalotis, Microtus ochrogaster and M. pennsylvanictis were taken in the same trapline with E. 7n. pallidus in Wind Cave Canyon.
A male taken near Minnekahta in mid-June 1968 was molting over the rump and dorsum; scattered patches of new hair are interspersed among the older pelage on the sides and venter of another male obtained at the same time and place. Two adult females captured under these circumstances showed no sign of molt ( still were in winter pelage ) . Molt was manifest over the shoulders and dorsum of a female collected in Wind Cave National Park on 30 August 1968.
Four placental scars were evident in the uterus of a female obtained on 13 June 1968, but three other females col- lected in the summer showed no signs of reproductive activity. The average length of the testes of five males collected in the summer was 7.4 (7-9).
Two males captured in Wind Cave National Park on 31 August and 1 Sep- tember 1968, were parasitized externally by ticks, Ixodes spinipalpis Hadwen and Nuttall. Although external and cranial measurements are similar, E. m. pallidus
differs from E. m. silvoticus in being paler over the upper parts, sides, and underside of tail. When compared with specimens from eight other localities in the Hills, individuals from near Minnekahta and Wind Cave National Park are signifi- cantly more pale in color tone ( see Table 8). Selected external and cranial meas- urements of three male and four female pallidus from the Black Hills are: total length, 203.0 ±5.66, 197.0 ±9.80; tail length, 89.7±3.53, 75.3± 12.53; hind foot length, 32.0±0.00, 32.0±0.82; ear length, 16.7±1.41, 18.4±1.89; weight, 52.9±3.96, 53.3±3.39; greatest length of skull, 32.7±0.18, 32.8±0.84; zygomatic breadth, 18.4 ±0.46, 18.2 ±0.30; brain- case breadth, 15.9±0.14, 15.6±0.41; ros- tral length, 11. 7 ±0.28, 11. 7 ±0.44; inter- orbital breadth, 7.5±0.00, 7.0±0.53; maxillary tooth-row length, 5.2 ±0.04, 5.2±0.17; cranial depth, 13.3 ±0.14, 13.2 ±0.48.
Specimens examined (15). — SOUTH DA- KOTA: Custer Countij: Beaver Creek Canyon, Wind Cave National Park, 4200 ft, 1; Elk Mountain Campground, Wind Cave National Park, 4200 ft, 1; Headquarters, Wind Cave National Park, 4100 ft, 2; Wind Cave Canyon, Wind Cave National Park, 4100 ft, 2; Wind Cave National Park, 4 (2 UMMZ, 2 WCNP). Fall River County: 0.5 mi S, 1.5 mi W Minne- kahta, 4200 ft, 5.
Eutamias minimus siK aticus White
Least Chipmunk
Eutamias minimus silvaticus White, 1952, Univ. Kansas Puhl., Mus. Nat. Hist, 5:259-262, 10 April (type locality, 3 mi NW Sundance, 5900 ft, Crook Co., Wyoming).
The subspecies silvaticus is endemic to the Black Hills. Eutamias minimus is represented by three morpliologically similar, but distinctively colored, races on the Northern Great Plains as follows: the dark sdvaticus of the conifer-clad Hills, the paler pallidus of surrounding foothills and wooded lowland escarp- ments and the extremely pallid caco- demus of the barren alkaline topography to the east in the Badlands National Monument and vicinity (Table 8). Least chipmunks are extremely common and
TURNER: MAMMALS OF THE BLACK HILLS
65
Table 8. — Geographic \arialion in coloration of the mid-dorsal pelage of adult Eutamias minivnis obtained in summer from the Northern Great Plains. Dashed lines separate populations that differ signifi- cantly.
Number and sex
of specimens
averaged
Hot*
o
c
a!
O
n3 CO
a o
C fuqn
PL, W) S
0)
« s
O <D Oj 3 W
150 (42 5, 108$ )
Mean SD
E. in. silvaticus. Black Hills of South Dakota
and Wyoming
28.4 46.4 28.8 24.8
±3.33 ±0.42 ±0.27 ±0.28
12 (4ci,8$)
Mean SD
8 (65,25)
Mean SD
6(35,35)
Mean SD
5(25,35)
Mean SD
2 (15,15)
Mean SD
E. m. paUidus, 2 mi N, 5 mi W Ludlow,
Harding Co., South Dakota
33.0 45.2 28.8 26.0
±3.39 ±0.40 ±0.26 ±0.22
E. 1)1. paUidus, 10 m S, 5 mi W Reva,
Harding Co., South Dakota
35.8 44.2 29.7 26.1
±6.90 ±0.25 ±0.24 ±0.27
E. m. paUidus, northern Campbell County,
Wyoming
36.4 45.7 28.8 25.5
±3.71 ±0.29 ±0.25 ±0.47
E. m. paUidus, 0.5 m S, 1.5 m W Minnekahta, Fall River Co., and Wind Cave
National Park, Custer Co., South Dakota
39.3 45.2 29.3 25.5
±3.82 ±0.25 ±0.11 ±0.20
E. m. paUidus, northern Siou.x County.
Nebraska
40.0 50.6 28.8 20.6
±2.12 ±0.35 ±0.29 ±0.33
5(15,45)
Mean SD
4 (35,15)
Mean SD
6(15,55)
Mean SD
E. m. cacodernus, 14 m N Long Valley,
Washabaugh Co., South Dakota
58.0 43.7 30.1 26.2
±4.05 ±0.25 ±0.16 ±0.13
E. m. cacodernus, 10 m N, 4 mi E Potato
Creek, Washabaugh Co., South Dakota
69.9 44.4 30.5 25.1
±9.42 ±0.24 ±0.14 ±0.10
E. m. cacodernus, 6 m S, 2 mi W Scenic,
Pennington Co., South Dakota
73.8 43.3 30.9 25.8
±5.90 ±0.11 ±0.13 ±0.13
widespread in all counties that comprise the Black Hills. Population densities seem to be much lower in the southern foothill zone and along the hogback ( see
account of E. m. pallichis). The species is most numerous in the forested uplands of the boreal cap and its presence has been noted even near the summit of
66
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Harney Peak (7240 feet) and atop Devils Tower. Eutamias minimus hibernates in the colder months. Individuals inhabit- ing peripheral foothills and lower eleva- tions probably hibernate for a shorter period ( specimens have been taken from 9 February to 20 November in these areas) than do inhabitants of the colder recesses of the mountainous Central Ba- sin. Least chipmunks were first reported from the Black Hills by Baird (1858: 299) under the name Tamias quadrivit- tatus.
Individuals captured in the present study either were shot or were taken in museum special snap-traps baited with rolled oats. Eutamias minimus appar- ently is ubiquitous, occurring in all habi- tats except extensive grasslands, which are inhabited instead by Spermophilus tridecemlineatus. In areas where border- ing woodland interdigitates with grass- land, and in campgrounds, these two small sciurids may be closely associated. Sparsely vegetated bluffs, rocky outcrops, and deteriorating abandoned buildings are environs shared with Neotoma cine- rea. Pine-clad slopes, spruce-filled can- yons, deciduous riparian woodland and brushy meadows that contain piles of logs are all inhabited by least chipmunks, which at one place or another have been obtained in association with Sorex cine- reiis, Peromijscus leucopus, P. manicu- latus, Reithrodontomys megalotis, Cleth- rionomijs gapperi, Microtus longicaudus, M. pemisylvajiicus and Zapus hudsonius. Eutamias minimus is most conspicuous in old burned areas that are regenerating with secondary vegetative growth, and in logged areas with abundant trimmings and fallen trees. For example, from 6 to 8 September 1968, 76 individuals (39 males and 37 females) were obtained in Roby Canyon along the western edge of the Black Hills; these were taken mainly in logged areas near Roby Springs and on mountain mahogany-dominated slopes near the mouth of the Canyon.
Bailey (1888:437) noted that chip- munks in the Black Hills feed largely upon seeds of wild roses, wild rye, snow-
berries, asters, chokecherries, and various grasses. A male shot in Ditch Creek Valley on 14 June 1965 was carrying a juniper berry in its mouth. Reforestation programs in the Hills may suffer a 50 to 75 percent annual seed loss due to chipmunks and other small rodents (Henshaw, 1910:551); a single chipmunk was observed visiting 38 seed spots in four minutes in one experimental plot, which suffered 70 percent loss in a three- day period (Silver, 1924:167). Baits of oatmeal mixed with strychnine and wa- ter, and wheat coated with hot tallow mixed with strychnine previously have been used to control chipmunk popula- tions in the Black Hills (Henshaw, loc. cit.).
White (1953b: 588-589) described the process of molt in E. m. silvaticus, based on study of a large series of specimens taken in several seasons of the year. I concur with his general analysis. Sum- mer molt of chipmunks in the Black Hills begins in mid-June and is com- pleted in about mid-August. Seasonal charge in color affects mainly the tints [tone and hue], whereas the overall pat- tern remains constant throughout the year, and is similar in both juvenal and adult pelages (J. A. Allen, 1890:49). Comparison of pelages of 53 adults taken in June ( 16 males and 37 females ) with 101 adults (28 males and 73 females) obtained in July indicates that the new fur of late summer reflects reds at a sig- nificantly greater intensity than does the older pelage of early summer. Measure- ments for the mid-dorsal region of speci- mens collected in June are followed by those from July: color tone, 29.5 ±5.57, 28.3 ±3.67; percent red reflectance, 44.7 ±0.43, 47.2 ±0.38; percent green reflect- ance, 29.7±0.25, 28.4±0.24; percent blue reflectance, 25.6±0.27, 24.5±0.26. Change in pelage is manifest either by distinct molt lines, or by interspersion of new hairs beneath the older fur. Most juvenile and subadult chipmunks ob- tained in the warmer months were in some stage of pelage replacement, indi- cating a gradual process as individuals
TURNER: MAMMALS OF THE BLACK HILLS
67
mature throughout the summer. Adults taken on the same day at the same lo- eahty may evidence different stages of molt, whereas others show no sign of molt. Pelage replacement may be de- la\ed in breeding females (Howell, 1929:28). Of 70 adult females obtained in the summer that showed signs of re- production (embryos, placental scars, or lactation ) , 80 percent manifested no sign of molt; the corresponding figure for 44 non-breeding adult females was 58 per- cent.
Of 54 adult females examined in June, 15 showed no signs of reproductive activity, 18 were lactating, 17 had an average of 5.1 (3-8) placental scars in the uteri, and four pregnant individuals carried 4.5 (3-6) embryos that were 13.7 (7-27) in crown-rump length. Twenty- four males captured in this period had testes that were 8.5 (4-17) in average length. Of fifty-seven adult females ex- amined in July, 27 were non-breeding, 20 were lactating, 10 evinced 5.0 (3-7) placental scars, and none was pregnant. The average testicular length of 17 males collected in July was 6.1 (4-10). None of eight adult females examined in August was pregnant and reproductive activity was not evident in two of these; however, two others were lactating and 5.5 (5-6) placental scars were present in the repro- ductive tracts of the remaining four fe- males of this series. The average testic- ular length of six males was 6.8 (6-8) in August.
Two females obtained west of Nemo on 29 June and 3 July 1967, and a male taken at Roby Springs on 7 September 1968 were parasitized externally by ticks, Dermacentor andersom Stiles. Other least chipmunks captured at Roby Springs on the same date harbored another kind of tick, Ixodes spinipalpis Hadwen and Nuttall, lice, Hoplopleum arboricola Kellogg and Ferris, fleas, MonopsyUiis eumolpi (Rothschild), and fur mites, Dermacanis hypudaei (Koch).
In considering geographic variation in E. minimus, cranial measurements were taken in the manner described by
White (1953a:566) and aging criteria also followed White ( 1953b : 587-588 ) . There were no significant differences in dimensions of adults and old subadults, and both age groups were consolidated when making taxonomic comparisons. In contrast to the findings of White (loc. cit.), females were significantly larger in total body length, weight, greatest length of skull, zygomatic breadth, length of rostrum, and length of maxillary tooth- row (although absolute differences are rather small for some of these means, minor variation in a parameter that is uniformly constant will result in statis- tically significant differences). There- fore, sexes were treated separately when making comparisons; secondary sexual variation in coloration was not apparent.
In the Black Hills, specimens from the Bear Lodge Mountains, and western areas in general, are more reddish than individuals from elsewhere; conversely, reflection readings obtained with green and blue filters are higher for eastern populations. For example, localities in the Hills were consolidated into five groups and resultant average percent re- flectance of reds for each group follows: Bear Lodge Mountains, 50.0 ±0.26; northwestern section, 48.0 ±0.32; south- western section, 46.7 ±0.24; southeastern section, 44.7±0.54; and northeastern sec- tion, 42.9±0.34. Individual localities within these sections also follow this general trend. A distinct pattern of morphological variation within the Hills was not discernible. Selected external and cranial measurements of 54 males and 129 females are: total length, 194.9 ±12.82, 201.0± 12.22; tail length, 81.0± 8.45, 83.3± 10.27; hind foot length, 31.0 ±1.61, 31.8±4.73; ear length, 16.3±1.29, 16.1±1.55; weight 46.8±3.89, 55.0± 6.97; greatest length of skull, 32.4 ±0.69, 32.6±0.48; zygomatic breadth, 18.4± 0.36, 18.6 ±0.39; breadth of braincase, 15.7 ±0.39, 15.7 ±0.34; rostral length, 11.4±0.30, 11.5±0.28; interorbital breadth, 7.0±0.32, 7.0±0.27; maxillary tooth-row length, 5.2±0.14, 5.3±0.15; cranial depth, 13.2±0.27, 13.2 ±0.23.
68
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Specimens examined (612). — SOUTH DA- KOTA: Lawrence County: Spearfish, 3 (SDSU); 3 mi S Speai-fish, 4200 ft, 1; White- wood, 2 (USD); Tinton, 2 mi N, 13 mi W Lead, 5900 ft, 30; 2 mi S Tinton, 6100 ft, 78; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 5800-6000 ft, 7; Little Spearfish Canyon, Sa- voy, 8 mi W Lead, 3 (USNM); 4 mi S, 7 mi W Cheyenne Crossing, 1; Deadwood, 19 (USNM); Strawberry Hill Campground, 3 mi S, 4 mi E Lead, 1 (SDMT); Dumont, 1 (USNM); Minnesota Ridge, 1 (SDSU); 2 mi W Nemo, 4700 ft, 22; Nemo, 4700 ft, 2 (1 SDSU); Blackfox Campground, 8 mi NW Rochford, 2 (NRW). Meade County: Vanocker Canyon, 3 mi S, 1 mi W Sturgis, 4600 ft, 1; Haven Dams, 6 mi S, 1.5 mi W Sturgis, 4600 ft, 5; 6 mi S, 1 mi W Sturgis, 4500 ft, 1. Pennington County: Rochford, 3 (UMMZ); South Canyon, 3-4 mi W Rapid City, 5 (AMNH); Dark Canyon, 2 mi S, 5 mi W Rapid City, 3800 ft, 9 (8 AMNH); Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 50; 3 mi N, 9 mi W Deerfield, 6500 ft, 5; 3 mi N, 7 mi W Deerfield, 6900 ft, 2; Deerfield Lake, 6400 ft, 1; Castle Creek, 6500 ft, 2 (UMMZ); 2 mi S, 1 mi W Pactola, 1 (SDSU); Redfern, 5 (USNM); Sheridan Lake, 1 (SDMT); Rockerville, 16 (2 NRW, 14 UMMZ); 3 mi S, 1 mi W Rocherville, 2; 20 mi N Elk Mountain, 6000 ft, 3 (USNM); 4 mi NW Hill City, 2 (UMMZ); Ditch Creek, 14 mi W Hill Citv, 6400 ft, 29; Clendale, near Hill City, 19 ( 15 AMNH, 4 FMNH); Hill Cit>-, 4 (1 NRW, 3 USNM); 2 mi S, 13 mi W Hill Citv, 6400 ft, 1; 4 mi S, 16 mi W Hill City, 1; 17 mi NW Custer, 1 (UMMZ); 16 mi NW Custer, 53 (UMMZ); Palmer Gulch, 3-4 mi SE Hill City, 5300-5400 ft, 20 (UMMZ); Horse Tliief Lake, 3 mi W Mount Rushmore National Monument, 5100 ft, 1; unspecified lo- cality, 5 (UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, 5300 ft, 5 (FMNH); 3.5 mi S, 0.5 mi W Keystone, 5000 ft, 15; 5.75 mi N, 5.75 mi E Custer, 5200 ft, 22; Roby Springs, 4 mi N, 22 mi W Custer, 5400 ft, 12; Roby Canyon, 2 mi N, 22 mi W Custer, 5200 ft, 4; Bull Springs, 2 mi N, 9 mi W Custer, 6500 ft, 8 (UMMZ); 0.9 mi W Custer, 1 (SDMT); Custer, 5300 ft, 13 (4 AMNH, 2 UMMZ, 7 USNM); Bismark Lake Camp- ground, 1 mi E Custer, 1 (SDMT); 4 mi SW Custer, 3; Squaw [=Grace Coolidge] Creek, 1 (AMNH); Custer State Park, 21 (UMMZ); Campbell's Ranch, Elk Mountain, 4800 ft, 5 (1 UMMZ, 4 USNM); Mayo, 1 (USD); 3 mi SW Pringle, 2; unspecified localitv, 5 (4 UMMZ, 1 USD).
WYOMING: Crook County: Devils Tower, 4 (USNM); Warren Peak, Bear Lodge Moun- tain, 6000 ft, 3 (USNM); 15 mi N Sundance, 5500 ft, 6; 15 mi ENE Sundance, 3825 ft, 1; 3 mi NW Sundance, 5900 ft, 17; 1 mi N Sun- dance, 1; Sundance, 8 (USNM). Weston
County: 1.5 mi E Buckhorn, 6150 ft, 19; 4 mi E Four Corners, 6; 9 mi N, 1 mi E Newcastle, 2; 6 mi N Newcastle, 1; Newcastle, 3 (USNM); SE Newcastle, 1 (USNM).
UNSPECIFIED LOCALITY. Black Hills, 4 (2 AMNH, 2 SDSU).
Additional records.— SOUTH DAKOTA: Pennington County: Rapid Citv (Howell, 1929: 57). WYOMING: Crook County: 2.5 mi N Sundance (Sutton and Nadler, 1969:526).
Marmota flaviventris dacota (Merriam)
Yellow-bellied Marmot
Arctomys dacota Merriam, 1899, N. Amer. Fauna, 218, 30 October (type locality. Black Hills, Custer, Custer Co., South Dakota).
M[armota]. fUaviventer]. dacota — A. H. Howell, 1914, Proc. Soc. Washington, 27:15, 2 Feb- ruary.
[M.] flaviventris dacota — A. H. Howell, 1915, N. Amer. Fauna, 37:7, 7 April.
Yellow-bellied marmots are distrib- uted in montane areas of the western United States, being confined to moun- tains, foothills and rocky canyons, and not occurring on the plains proper. Baird (1858:344) first noted the occurrence of this marmot in the Black Hills, as Arc- tomys flaviventer. The subspecies dacota is named from, and restricted to, the Black Hills of South Dakota and Wvom- ing, where it inhabits rocky hillsides, crevices in bluffs, rockpiles in meadows, or abides beneath deserted buildings. Altitudinal distribution is from the bo- real cap of the Hills, down to about 4000 feet. Specimens of this species have been recorded from all counties that comprise the study area. Yellow-bellied marmots hibernate during the colder months; the earliest and latest seasonal records from the Hills known to me are 12 April and 22 August, respectively.
Distribution of Marmota flaviventris in the Hills coincides with the occurrence of rocky outcroppings or other high van- tage points from which these animals can sun themselves and survey the sur- rounding countryside. These vantage points are closely associated with mon- tane meadows, pastures, and fields of alfalfa and clover. Vernon Bailey found flowers, leaves, and green seeds of vari- ous plants, including milk- vetch (As-
TURNER: MAMMALS OF THE BLACK HILLS
69
truiialu.s hi.siilcatus) and stonecrop (Se- (lui)i doiiiilasii) in the stomachs of mar- mots from the Black Hills (Howell, 1915:13).
Specimens from Ditch Creek Valley, Beaver Creek \'alley, and Little Spear- fish Canyon generally were taken in habi- tats as described above. An adult female and seven juveniles (four females and three males ) obtained southwest of Key- stone were shot at the entrance of an old lead-zinc mine and in adjacent wood- piles. The mine was on a hillside that was clad in pine and bur oak; well-used pathways were evident leading from the mine to a valley meadow of bluegrass and clover below.
A field party from The University of Kansas dug an adult female and six young males out of a den in Renter Can- yon on 4 July 1947. The burrow was in a hillside that faced north and the tunnel ran obliquely upward about 10 degrees from the horizontal for approximately 12 feet, leveled off and ran at a right angle for another six feet; no nest was located.
A graduate student from the Museum of Natural History, Paul B. Robertson, spent several days observing a colony of yellow-bellied marmots (five adults and seven juveniles) at a locality 4 mi W Nemo, Lawrence County. The colony occupied an area of about 500 square yards that comprised an old homestead. Deserted buildings, rock piles, and wood- piles were situated in a meadow and a pasture. Activity of the marmots began about 6:45 hrs (MDT) on sunny morn- ings, with more restricted activity on cloudy days. After an initial period of vigilance, marmots spent most of the morning grazing on the meadow. Move- ment of adults was much more conserva- tive than that of the young; while adults remained alert, young romped freely, stopping often to preen or graze. When feeding in tall grass, marmots raised up on their haunches much more frequently than in short grass, and feeding was more sporadic. In late morning and early afternoon, grazing activity declined and sunning behavior increased. On one
occasion, the colony sensed the approach of a man while he was yet one-quarter of a mile distant. After a series of whistles from the adults, young animals retreated toward the burrow systems, while the older marmots remained alert until the man passed from sight. Graz- ing activity increased once again about 16:30 hrs (MDT) and continued until near dusk.
Two pregnant females obtained south- east of Hill City on 12 April 1946 con- tained five and six embryos that were 16 and 9 in crown-rump length, whereas a female from near Keystone (22 June 1967) and another from Little Spearfish Canyon (1 August 1967) evidenced eight and two placental scars, respec- tively. Lactating females were taken on 13 and 22 June 1967, and on 8 July 1965; young marmots were common from early July through August. Testicular length of an adult male collected on 12 June 1965 was 12, and testes of six subadult males captured early in August 1967 had an average length of 8.2 (7-10).
The venter, feet, tail, and underfur of juveniles from near Keystone are a dull dark brown, with a pale brown- or beige-tipped guard hairs over the dorsum and sides, in contrast to the brightly col- ored reddish hairs of adults. Change from subadult to adult pelage and an- nual molt in adults occurs from mid- June through July. Replacement of the old pelage is most noticeable over the mid- dorsum, sides, and rump; replacement of the mantle (a cloak of long hairs covering the shoulders and anterior third of the dorsum) is more difficult to ob- serve but on occasion new hairs are evident. The adult female from near Keystone seemed to have a mange-like disease and lacked hair entirely over the mid-dorsum.
Marmota faviventris dacota differs from M. /. luteola of south-central Wyo- ming (Converse, Albany, and Carbon counties ) , in the greater contrast in color between the mantle and mid-dorsal pel- age (Table 9) and brighter color of the mantle of the former. Howell (1915:50)
70
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Table 9. — Geographic variation in coloration (mean and standard devia- tion) of the mantle and mid-dorsal pelage of adult Marmota flaviventris obtained in summer from the Northern Great Plains.
Number and sex
of specimens
averaged
s o
|
<a |
<D |
<D |
|
|
o |
O |
O |
|
|
5 |
4-' |
a |
c « |
|
Si y |
o |
c o |
Oj cj |
|
O 0^ |
o |
OJ 0* |
O 0) 1< |
|
111 |
13 (4^,92)
Mantle Dorsum
8 (2(5,69) Mantle Dorsum
10 (2(5,89 ) Mantle Dorsimi
M. /. dacota, Black Hills of South Dakota
and Wyoming
51.4+8.59 54.9±0.23 25.3±0.16 19.8±0.10
36.3±5.40 53.1±0.75 23.9±0.39 23.0±0.42
M. /. luteola, south-central Wyoming 39.9±6.89 54.2±0.30 25.9±0.29 32.8±5.59 56.8±0.59 23.1
:0.36
19.9+0.24 20.1±0.26
50.3: 34.3:
M. f. nosophora, western Wyoming :11.66 50.3±0.36 26.1 ±0.34 23.6±0.23 :5.44 51.3±0.53 25.3±0.14 23.4±0.48
noted that dacota is the brightest of all races of the species, with red and yellow shades being most pronounced and blacks and browns reduced to a mini- mum. Indeed, along with that of Tamia- sciunis hudsonicus, pelage of yellow- bellied marmots reflects reds at greater intensities than that of other species examined from the Black Hills. Marmota flaviventris luteola also lacks the blackish muzzle of M. f. dacota, and has some- what smaller average dimensions; total btxly length (577.5 it 36.45) and inter- orbital breadth (17.7±1.41) of two male and si.x female luteola differ significantly from those of six male and 10 female dacota (620.7±47.06 and 20.3 + 2.06, re- spectively ) . Intergradation between these two subspecies is evident in speci- mens from the Laramie Mountains, where individuals are more reddish in color and the total body length and interorbital breadth are greater than in typical Ititeola.
Marmota flaviventris nosophora of western Wyoming ( Park, Big Horn, Wa- shakie, Sublette, Fremont and Natrona counties) is similar to dacota, but has slightly smaller average dimensions; the upper parts differ significantly in reflec- tion of reds, are slightly darker in tone, and are more mixed with black; the
mantle of nosophora reflects blues at sig- nificantly greater intensities than that of either dacota or hiteola (Table 9). Speci- mens similar to each of the three sub- species can be found in south-central Wyoming due to intergradation and the high degree of individual variation in yellow-bellied marmots of that area (Long, 1965:568).
Significant secondary sexual differ- ences in color and size of M. /. dacota are not apparent, nor is a pattern of variation within the Black Hills readily discernible. Selected external and cranial measurements of six males and 10 fe- males from the Hills are: total length, 620.7±47.06; tail length, 178.7+24.52; liind foot length, 79.9+3.54; ear length, 29.3+6.32; weight (kilograms), 3.0 + 0.83; greatest length of .skull, 84.3+3.73; zygomatic breadth, 55.9+2.73; rostral lengtli, 33.8 + 2.04; interorbital breadth, 20.3 + 2.06; postorbital breadth, 16.9+ 0.93; mastoid breadth, 40.6 + 1.99; and length of maxillary tooth-row, 20.9+0.61.
Specimens examined (73). — SOUTH DA- KOTA: Lawrence County: Tinton, 2 mi N, 13 mi W Lead, 5900 ft, 4; Little Spearfish Canyon, Savoy, 5 (USNM); Little Spearfi.sh Canyon, 1 mi S, 9 mi W Lead, 5400 ft, 4; Little Spearfi.sh Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 4; 3 mi W Nemo, 4800 ft, 1; 1 mi E Nemo, 4700 ft, 1. Meade County: Vanocker
TURNER: MAMMALS OF THE BLACK HILLS
71
Canyon, 6 mi S, 1 mi W Sturgis, 4500 ft, 2. Pennington County: Beaver Creek Valley, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 2; Dark Canyon, 2 mi S, 5 mi W Rapid City, 1 (AMNH); Sheridan Lake, 6 mi NNW Key- stone, 1; 5.4 mi E Mount Rushmore National Monument, 1 (SDMT); Ditch Creek Valley, 14 mi W Hill Citv, 6400 ft, 7; Tigerville, 1 (USNM); Palmer Gulch, 3 mi SE Hill City, 5300 ft, 2 (UMMZ). Custer Counti/: Palmer Gulch, 8 mi SE Hill Citv, 5400 ft, 1 "(FMNH); 3.3 mi S, 0.5 mi W Kevstone, 5000 ft, 8; Custer, 13 (7 USNM, 6 AMNH); Doran's Ranch, 3 mi E Custer, 1 (UMMZ). Fall River County: 5.5 mi E Minnekahta, 4000 ft, 1.
WTOMING: Crook County: Renter Can- yon, 1 mi S Warren Peak, 6000 ft, 8; Bear Lodge Mountains, 1 (USNM). Weston County: 0.5 mi E Buckhorn, 6100 ft, 1; 1.5 mi E Buckhorn, 6150 ft, 1.
UNSPECIFIED LOCALITY: Black Hills, 2 (1 USNM, 1 FMNH).
Spermophilus tridecemlineatus pallidas
J. A. Allen
Thirteen-lined Ground Squirrel
Spermophilus tridecem-lineatus var. pallidus J. A. Allen, 1874, Proc. Boston Soc. Nat. Hist., 16:291, June (nomen nudum).
Spermophilus tridecemlineatus var. pallidus J. A. Allen, 1877, in Coues and Allen, Bull. U. S. Geol. Surv. Territories, 11:872, Au- gust (type locality restricted to the mouth of the Yellowstone Ri\er, McKenzie Co., North Dakota, by A. H. Howell, N. Amer. Fauna, 56:122, footnote, 18 May 1938).
Spermophilus tridecemlineatus olivaceus J. A. Allen, 1895, Bull. Amer. Nat. Hist., 7:337, 8 November (type locality. Black Hills, Custer, Custer Co., South Dakota).
The thirteen-lined ground squirrel in- habits dry grasslands of the central United States, and is locally common within the Black Hills in areas of short grass such as moderately grazed pastures, mowed borders of roadways and camp- grounds, and upland meadows. This spermophile ranges through the foothill transition zone up to 6500 feet, but is more common below 5500 feet. Spermo- philus tridecemlineatus hibernates in the study area from mid-September until early April; the earliest and latest sea- sonal records as evidenced by the speci- mens examined are 8 April and 7 Sep- tember, respectively.
Grinnell (1875:82) observed large
numbers of this species on the surround- ing plains, but saw none while in the black Hills; S. tridecemlineatus was first noted therein by Bailey (1893:45). Ground sciuirrels often arc^ observed dur- ing the day along roadsides and in the parkways of campgrounds. Traps placed in these areas took five specimens in Ditch Creek Valley in mid-June 1965; another was shot as it ran from its bur- row after water was poured into the tunnel system. In Wind Cave National Park, two individuals were taken on 27 August 1968 along a fence that served as a corral for bison. The enclosed area was closely grazed, whereas exterior to the fence, natural prairie grasses and forbs were about knee-high. J. A. King obtained several S. tridecemlineatus in early April in an upland meadow that was being grazed by horses; one individ- ual was carrying unidentified seeds in its cheeks. Fifty percent of the stomach contents of a female taken at Custer in July 1893 was composed of grasshoppers and other insects, the remainder being seeds and vegetable matter (Bailey, 1893).
Reithrodontomys megalotis, Peromy- scus maniculatus, Microtus ochrogaster, and Eutamias minimus often were taken in the same trapline with the thirteen- lined ground squirrel. Long (1965:581) stated that he had seen Spermophilus tridecemlineatus and Eutamias minimus "so closely associated in Custer County, South Dakota, that the latter could be chased into the burrows of the former."
Four of eight females (for which reproductive data are available) taken from 12 June to 1 July, were pregnant, averaging 7 (3-9) embryos that were 17 (6-27) in crown-rump length; another individual was lactating. Testes of five males obtained in the same period were 10.4 (5-16) in length. Juveniles first ap- peared above ground in early July, and were common by mid-summer.
Seven individuals obtained in mid- June west of Hill City were molting con- spicuously over both sides and the dor- sum. A female from Wind Cave National
72
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Park harbored several ticks, Ixodes sculp- tus Neumann, and fleas, Thrassis joins ( Jordan ) .
J. A. Allen (1895b: 337) described a new subspecies, Spermophilus tridecem- lineatus olwaceus from the Black Hills; he indicated that it differed from the nearby pallidus "in its much darker ground color and the olivaceus creamy white tint of the light stripes and spots." A. H. Howell (1938:113) placed oliva- ceus in the synonomy of pallidus, stating: "The type series of olivaceus from Custer, South Dakota, lias been compared with a large series of typical pallidus and is found to agree closely with it." Long ( 1965:581 ) reinstated olivaceus as a sub- species of S. tridecemlineati4S, indicating tliat in addition to the difference in col- oration noted by Allen, olivaceus also is characterized by shorter nasals, larger auditory bullae, and longer external ears than pallidus.
I have compared a large series of specimens from the Black Hills with typical pallidus from northwestern (Harding County) and southwestern ( Bennett County ) South Dakota, north- and western Nebraska (Sioux,
ern
Cherry, Keya Paha, and Cheyenne coun- ties), northeastern Wyoming (Campbell, and Weston counties), and southeastern Wyoming (Albany, Goshen, and Laramie counties ) . Morphologically, spcrmo- philes from the Black Hills fall within the range of variation of the comparative material and do not differ significantly from pallidus in anv dimension (Table 10).
Ground color and coloration of stripes and spots on individuals of S. tridecem- lineatus vary greatly, and most possible color combinations are present when a large series of specimens is examined. On the average, mid-dorsal pelage of thirteen-lined ground squirrels from the
Table 10. — Geographic variation in selected external and cranial measurements of adult Spermophilus tridecemlincatus from the Northern Great Plains.
|
Number and sex of specimens averaged |
Total length |
Ear length |
Greatest length of skull Zygomatic breadth Rostral length Length of maxillary tooth-row^ |
Skull depth |
Greatest diameter of audi- tory bullae |
|
21 (105,119) |
Black Hills, South Dakota and Wyoming |
||||
|
Mean |
232.1 |
9.1 |
38.5 22.5 14.8 7.2 |
16.7 |
8.4 |
|
SD |
±7.44 |
±1.64 |
±1.08 ±0.86 ±0.63 ±0.33 |
±0.39 |
±0.33 |
|
4 (U,35) |
Harding County, South Dakota |
||||
|
Mean |
241.2 |
8.0 |
38.3 23.7 15.0 7.2 |
16.4 |
8.2 |
|
SD |
±13.74 |
±1.41 |
±0.60 ±0.65 ±0.47 ±0.31 |
±0.07 |
±0.32 |
|
3(1,5,2$) |
Bennett County, South Dakota |
||||
|
Mean |
242.7 |
9.0 |
39.5 23.3 15.5 7.3 |
16.5 |
8.5 |
|
SD |
±11.68 |
±1.00 |
±0.64 ±0.12 ±0.35 ±0.29 |
±0.26 |
±0.00 |
|
7(25,5?) |
Northeastern Wyoming ( see text ) |
||||
|
Mean |
223.7 |
9.7 |
37.6 22.3 14.3 6.9 |
15.8 |
8.1 |
|
SD |
±19.39 |
±1.03 |
±1.38 ±0.94 ±0.73 ±0.28 |
±0.61 |
±0.12 |
|
11( 55,69) |
Southeastern Wyoming (see text) |
||||
|
Mean |
228.9 |
7.2 |
38.1 22.5 15.1 7.0 |
16.4 |
7.9 |
|
SD |
±16.33 |
±1.32 |
±1.27 ±0.79 ±0.68 ±0.18 |
±0.61 |
±0.33 |
|
5(25,39) |
] |
Vorthern and western Nebraska ( see text ) |
|||
|
Mean |
246.0 |
9.4 |
38.9 23.6 15.3 7.4 |
16.7 |
8.4 |
|
SD |
±6.68 |
±1.52 |
±1.91 ±1.07 ±0.74 ±0.28 |
±0.36 |
±0.17 |
TURNER: MAMMALS OF THE BLACK HILLS
•73
Hills is dark in tone, high in reflectance of reds, and low in reflectance of blues (Table 11). However, these individuals readily fall within the range of variation of populations from Bennett County and from parts of W'Noming, and do not differ significantly from them. Thus, I must concur with Howell and place S. t. oliva- ceus in the synonomy of S. t. pallidiis.
Significant secondary sexual variation was not e\'ident. Although males aver- aged somewhat larger in se\'eral cranial dimensions, females were slightly larger in total length and length of tail. Addi- tional measurements (not listed on Ta- ble 10) of 31 S. t. pallidus from the Black Hills are: tail length, 75.3 ±7. 16; hind foot length, 31.5±1.40; weight, lU.Sib 22.73; and postorbital breadth, 11. 1± 0.57.
Specimens examined (102). — SOUTH DA- KOTA: Lawrence Countij: L.5 mi S, 0.5 mi E
Spearfish, 3800 ft, 1; Crowfeet [=Crow Peak?], Black Hills, 1 (AMNH). Meade County: Black Hawk, 1 (AMNH). Penninpton County: 0.5 mi N Silver City, 1 (SDMT); Rapid City, 2 (NRW); Beaver Creek Valley, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 1; Ditch Creek Val- ley, 14 mi W Hill City, 6400 ft, 6; 2 mi S, 13 mi W Hill City, 1; Pactola, 1 (USNM); Castle Creek, 6500 ft, 2 (UMMZ); Diamond S Ranch, near Rapid City, 2 (UMMZ); Palmer Gulch, 3-4 mi SE Hill City, 53-5400 ft, 7 (UMMZ); unspeciBed locality, 1 (UMMZ). Custer County: Palmer Gulch. 8 mi SE Hill City, 5300 ft, 4 (FMNH); Harney National Forest, 10 (UMMZ); Custer, 25 (1 UMMZ, 18 AMNH, 4 USNM, 2 FMNH); Bull Springs, 2 mi N, 9 mi W Custer, 10 (USNM); Elk Mountain, 1 (USNM); Buffalo Corral, Wind Cave National Park, 4300 ft, 2; Wind Cave National Park, 2 (WCNP).
WYOMING: Crook County: Bear Lodge Mountains, 5200 ft, 1 (USNM); 15 mi N Sun- dance, 5500 ft, 3; 1.5 mi NW Sundance, 5000 ft, 6; 1 mi N Sundance, 1; Sundance, 5 (USNM). Weston County: 1.5 mi E Buck- horn, 6150 ft, 4; Newcastle, 1 (USNM).
Table 11. — Geographic variation in coloration of the mid-dorsal
pelage of adult Spermophilus tridecemlineatus obtained in summer
from the Northern Great Plains.
Numlier and sex
of specimens
averaged
fl o H o o
a
111
O
C 03
O
a o
c
PL, bc S!
a
o
c
o
pm ^ S
21 (10c?, 112) Mean SD
Black Hills, South Dakota and Wyoming
33.8 47.3 28.9 23.8
±8.30 ±0.17 ±0.10 ±0.09
4 ( 1 ,5 , 3 9 )
Mean
SD
Harding County, South Dakota
45.0 45.3 30.1 24.6
±6.89 ±0.22 ±0.11 ±0.15
3 (U,29)
Mean
SD
Bennett County, South Dakota
32.8 47.2 28.4 24.4
±4.65 ±0.32 ±0.12 ±0.19
7(25,59)
Mean
SD
Northeastern Wyoming (see text)
41.9 45.4 30.2 24.4
±4.74 ±0.14 ±0.17 ±0.09
11 (56,69) Mean SD
Southeastern Wyoming ( see text )
47.4 45.6 29.2 25.2
±5.36 ±0.16 ±0.18 ±0.14
5(2<?,39)
Mean
SD
Northern and western Nebraska (see text) 49.5 45.5 29.7 24.8
±6.09 ±0.17 ±0.11 ±0.07
74
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Cynomys ludovicianus ludovicianus
(Ord)
Black-tailed Prairie Dog
Arctomys ludoviciana Ord, 1815, in Guthrie, A new geogr., hist., coml. grammar . . ., Philadelphia, Amer. ed. 2, 2:292, descrip- tion on page 302 ( type locality restricted to the "Tower," sec. 10, T. 34 N, R. 10 W, Bovd Co., Nebraska, by Jones, Univ. Kan- sas' Publ., Mus. Nat. Hist., 16:140, 1 Oc- tober 1964).
Cynomys hiclovicianus — Baird, 1858, Mammals, in Repts. Expl. and Surw . ., 8(1):331, 14 July.
The black-tailed prairie dog is a char- acteristic inhabitant of the short-grass plains of central United States. F. V. Hayden (1862:145) first noted this spe- cies in the Hills region: "The largest one [prairie dog town] I have seen is near the Black Hills, north of the Big Shey- enne river [Cheyenne River]. This vil- lage, though sometimes interrupted by high ridges or hills is connected, and covers an area of over fifty square miles." The Shirttail Canyon prairie dog town in the southwest corner of Wind Cave National Park was studied extensivelv by J. A. King (1955). C. B. Koford ( 1958 ) also studied the Shirttail Canyon colony, as well as the prairie dog town in Devils Tower National Monument. These two reports were consulted frequently in preparing the present account. Bailey (1888:440-441) described a prairie dog colony that occupied about 40 acres at a place 2 mi E Rapid City.
The Devils Tower prairie dog town comprises about 40 acres between the Belle Fourche River and the pine-clad hills of the National Monument. The town is at least 75 years old; in June 1894, Vernon Bailey noted "a good sized dog town occupies a level section in the valley at base of tower" (USBS files). Counts made in 1947 and 1955 indicated that the population was fairly stable at about 750 prairie dogs. In the area where Cynomys are fed by tourists, the number of burrows reaches a maximum density of 100 per acre.
The Shirttail Canyon prairie dog town occupies about 75 acres, covering
part of the canyon floor and adjacent ter- races. In 1950, King counted 614 bur- rows on an 11.7 acre sample plot of this town (52.5 per acre). Wind Cave Na- tional Park supports seven naturally es- tablished prairie dog towns and three new ones that were initiated in the north- westernmost corner of the Park in 1965. Cynomys ludovicianus occupies about 655 acres, or three percent of the non- forested portion (21,449 acres) of the Park. Total acreage inhabited by prairie dogs increased by 90 acres from 1963 to 1967.
This species is active the year around, although activity decreases somewhat during the colder months and on cloudy days. Both King and Koford found that males lose more weight than females during winter, possibly indicating that males are more active at that time. Bur- rows are constructed on floors of can- yons and on adjacent terraces, where fine-textured chernozems and chestnut soils enable prairie dogs to excavate deep, continuous passages; at Devils Tower, many burrows were located in fine sand of the Belle Fourche River floodplain. Barriers to expansion of prairie dog towns seem to be unsuitable soils of adjacent slopes, heavy cover, and rough terrain.
Earthen mounds thrown up around the entrances to burrow systems favor establishment of pioneer forbs such as bigbract verbena (Verbena hracteata), stickweed (Lappttla redowskii), dog- weed {Dyssodia papposa), and false pennyroyal (Hedeoma hispida). Koford analyzed the vegetation along a transect through the Shirttail Canyon colony and found that the center of the town sup- ported 67 percent buffalo grass, 24 per- cent tumblegrass (Schedonnardus panic- ulatus), six percent blue grama, and three percent western wheat grass. In the peripheral area, blue grama and west- ern wheat grass increased in abundance and the town border was dominated by Japanese brome grass (Bromus japoni- cus) . W^ithin the confines of a town, prairie dogs usually cut down tall plants
TURNER: MAMMALS OF THE BLACK HILLS
75
that otherwise would obstruct a clear field of observation; examples of such plants are woolly mullein, Russian thistle (Salsola kali), snowberry (Symphori- carpos occidentaJis), silver sagebrush, and brome grass. Other species sucli as snow-on-the-mountain (Euphorbia mar- ginata), clammyweed (Polanisia graveo- lens ) and \'arious mustards are left erect, presumably due to a bitter taste, presence of thorns, a tough integument, or other protective devices of the plants. In addi- tion to the numerous plant species listed by King ( 1955:8 and 9), Koford included scarlet globemallow, cutleaf nightshade (Solanum triforum), pepperweed (Le- pidiiim densifiorum) and lamb's-quarters as important foods of Cynomys liidovi- ciamis in the Black Hills region.
King reported that the Shirttail Can- yon colony was divided into fairly stable social groups of several members, each group occupying a particular territory of approximately one acre in size and each consisting of an adult male with a few females and their young. Even though a mother prairie dog defends the area of her nest burrow, Koford thought it was improbable that this aggressive- ness limits the number of families; for example, 11 nest burrows were found in an area of only 1.5 acres in Shirttail Canyon.
Expansion of a dog town presumably occurs when the population density in- creases shortly after the pups emerge from below ground. King noted that emergence of pups in May raised the average population density from four to 15 prairie dogs per acre. In contrast to the behavior of most rodents, adult prairie dogs move into a new area, from which the young are repulsed. The long- est movement recorded by King was 800 feet beyond the edge of the colony. Search for food may be another cause of colony expansion.
In early July 1968, I obsei-ved an adult badger and two young excavating a prairie dog burrow in Wind Cave Na- tional Park earh' in the morning. Sev- eral burrows in Shirttail Canyon also
were dug out by badgers, and coyotes are a fairly common sight in towns early in the morning. Although no actual sightings have occurred since 1953, char- acteristic trenches of the black-footed ferret were found in the National Park in 1969. King reported that a prairie dog was killed by a golden eagle in the winter of 1955; I observed an eagle swoop down upon an adult prairie dog in the summer of 1968, but the prairie dog escaped into its burrow. Remains of a young C. ludoviciamis were found in pellets of a great horned owl that was roosting in Wind Cave Canyon in July 1968.
Mating (occurs in February and March and two to 10 young are born in early April after a gestation period of 30 to 35 days. The pups emerge from under ground in mid-May. Evidently, only one litter is produced per year. King found no evidence of reproduction among yearlings and noted that mor- tality was high in the young during the first (36%) and second (22%) years.
King noted that fleas (Opisocrostis hirsutus), mites ( Africholaelaps f^Ias- f!,owi), and ticks (Ixodes sp.) parasitize prairie dogs externally in Shirttail Can- yon.
Specimens examined (29). — SOUTH DA- KOTA: Pennington Cotmit/: 0.5 mi E Rapid City, 1 (SDSU); Rapid City, 11 (USNM). Custer County: McAdam's Ranch, 3 mi NW Wind Cave, 1 (WCNP); Wind Cave National Park, 5 (2 UMMZ, 2 UW); Elk Mountain, 5 (UMMZ); 1 mi NE Wind Cave, Wind Cave National Park, 2 (WCNP).
WYOMING: Weston County. Newcastle, 4 (USNM).
Additional records.— SOVTYi DAKOTA: Meade County: Stnrgis (USES files). Custer County: Pringle (USES files); Horseshoe Eend, 12 miNE Elk Mountain (USES files).
Sciurus niger rufiventer
E. Geoffroy St.-Hilaire
Fox Squirrel
Sciurus rufiventer fi. Geofi"roy St.-Hilaire, 1803, Catalogue des mammiferes du Museum Na- tional d'Histoire Naturelle, Paris, p. 176 (type locality restricted to Mississippi Val- ley, probably between southern Illinois and
76
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
central Tennessee, Ijy Osgood, Proc. Biol. Soc. Washington, 20:44, 18 April 1907).
Sciurus niger rufiventer — Osgood, 1907, Proc. Biol. Soc. Washington, 20:44, 18 April.
The fox squirrel is a eommon resident of timbered areas and urban communi- ties of eastern and central South Dakota, but has been reported in Wyoming only from Cheyenne and Laramie (Long, 1965:598) \ind from the Black Hills (Maxwell and Brown, 1968:155). The distribution of this species in western South Dakota is essentially dendritic along the major river systems; for ex- ample, throughout the valley of the White River. In the Hills, the distribu- tion of Sciurus niger characteristically follows that of the bur oak and other deciduous elements, particularly on many slopes that lead up from stream beds in the northern Hills and the Bear Lodge Mountains, and in wooded ra- vines of the foothill zone.
Spread of the fox squirrel in Nebraska (Jones, 1964:146) and North Dakota (Hibbard, 1957:527) has been well doc- umented. This species probably would not occur in the study area were it not for the occurrence of gallery forests along river systems across the plains that served as avenues of dispersal and for introduction by man. I know of no in- troductions into the Black Hills proper, but fox squirrels are known to have been introduced in some adjacent areas (northwestern Nebraska, for example; see Jones, 1964).
Hayden (1856:79) indicated that the nothernmost limit of the fox squirrel in the plains region was at the mouth of Running Water [Niobrara Ri\'cr] on the Missouri, but later ( 1859:708) noted that this species reached the limits of its range at the mouth of the White River. I have observed these sciurids in resi- dential areas in Spearfish, Rapid City, and Hot Springs, and I have heard the distinctive bark of fox squirrel in Beaver Creek Canyon in Wind Cave National Park. Residents report this squirrel as being quite common along wooded streams east and north of Spearfish.
Specimens examined (5). — SOUTH DA- KOTA: Pennington County: Rapid Cit>', 3 (1 SDMT, 2 NRW); 0.75 mi SW Rapid City, 1 (SDSU).
WYOMING: Crook County: Sand Creek, SBeulah, 1 (UW).
Tamiasciurus hudsonicus dakotensis
(J. A. Allen)
Red Squirrel
Sciurus hudsonicus dakotensis J. A. Allen, 1894, Bull. Amer. Mus. Nat. Hist., 6:325, 7 No- \'ember (type locality, Squaw [=Crace Coolidge] Creek, Custer Co., South Da- kota).
Tamiasciurus luulsonicus dakotensis — Hayman and Holt, 1940, in Ellerman, The families and genera of living rodents, British Mus., 1:346, 8 June.
In the northern plains region, Tamia- sciurus hudsonicus inhabits montane for- ests and several scattered conifer-clad escarpments. The species is commonest on the boreal cap of the northern Black Hills where both white spruce and pon- derosa pine are predominant; the former conifer is scarce in the drier southern woodlands, and accordingly, population densities of Tamiasciurus are much lower in these areas. The red squirrel, or chickaree, is found in the Hills at elevations above 3800 feet, but is most numerous above 5200 feet.
Hayden (1859:708) first reported T. hudsonicus from the Black Hills, and J. A. Allen (1894a: 325) later described dakotensis is an endemic subspecies ( see Hall and Kelson, 1959:400, map 257). However, dakotensis is not restricted to the Hills, but occurs also in several out- lying areas. For example, Vernon Bailey captured a female at Belle Fourche, Butte Co., South Dakota, on 21 October 1887. B. W. Evermann obtained a male at Edgemont, Fall River Co., South Da- kota, on 7 October 1892, and V. H. Caha- lane took a male 5 mi S Hcrmosa, Custer Co., South Dakota, on 12 December 1934. J. A. Allen (1877:689) reported a speci- men from Bear Buttes, Meade Co., South Dakota. W. J. Hoffman (1877:97) noted that this species "sometimes is found in oak groxes eight miles west of the Post [Grand River Militaiy Post, Corson Co.,
TURNER: MAMMALS OF THE BLACK HILLS
77
South Dakota]," but this report is open to question. Swenk (1908:81) and Visher (1914:88) noted the former distribution of red squirrels in nortliern Nebraska and Harding County, South Dakota, respec- tively; however, Jones (1964:337-338) did not admit Taniiosciums to the fauna of Nebraska, and Andersen and Jones (1971:375) reported that the species evi- dently does not now occur in Harding Count)', although they did record a speci- men from the Long Pine Hills in adjacent Carter County, Montana, as dacotensis. Specimens of T. Jiudsonicus tentatively assigned to dakotensis have been re- ported from southeastern Montana and the northern Laramie Mountains in Wy- oming (Hoffman and Jones, 1970:374, Fig. 7).
This sciurid is active in the Hills the year around. Although this species does not hibernate, it may undergo periods of quiescence during prolonged inclement weather. A few individuals can be ob- served in summer at most hours of the dav, but maximal activity occurs between 7:00 and 9:30 hrs (MDT) and from 16:30 hrs to dusk. Even though popula- tion densities vary from area to area, the red squirrel is ubiquitous in wooded en- virons; pine-clad slopes, spruce-filled canyons, and oak-bordered drainage sys- tems all are inhabited. In the latter habitat, Tamiasciurus is ecologically sym- patric with Sciunis niger to an unknown extent. The fox squirrel is far less com- mon and its distribution in the Hills closely approximates that of bur oak and other deciduous elements.
While with a Museum of Natural History field party in 1947, R. B. Finley studied several external nests (located on branches of trees, rather than in a den cavity) of red squirrels in Crook County, Wyoming. He located three ponderosa pine nest trees on a 30"" slope with a southeastern exposure; the trees were from eight to 17 inches in diameter, and 30 to 60 feet in height. Nests were built 22.3 (18 to 25) feet above the ground and either were placed adjacent to the trunk, or rested upon a horizontal
limb (two to four inches in diameter) within two feet of the trunk. The loosely constructed nests were roughly oval in shape, measuring 15 (13 to 17) inches in length, 11.3 (9 to 13) inches in width, and 7.3 (6 to 8) inches in depth. In- teriors of the nests were shapeless, un- lined spaces, lacking a well-defined nest chamber. These were temporary or sum- mer nests, but characteristic winter nests are tightly constructed of twigs, lichens, mosses and other plant materials, and frequently were observed among the dense foliage of white spruce in the northern Black Hills.
Materials used in construction of sum- mer nests were varied, consisting of news- paper scraps (probably from a nearby abandoned Civilian Conservation Corps Camp ) , shredded cloth and bark, tufts of fur, and particles of lichens and dried graminoid plants. Another member of the field party, E. P. Marks, examined two of the above nests and found 18 dipteran larvae, three dipteran pupae, five flea larvae, two unidentified fleas, one pseudoscorpian and eight collembo- lans therein.
Tamiasciurus in the Black Hills seems to utilize cavities of trees more frequently than outside nests, especially when rear- ing young. On 27 June 1967, a male was shot from a den tree in Vanocker Can- yon, Meade County. The hollow pine tree was 30 feet in height and about 1.5 feet in diameter; the nest entrance was 2.5 inches in diameter and located near the top of the tree. After the den tree was pushed over, it was split open for a better view of the nest. Within the hollow was not one, but a sequential series of nests, graduated in age from bottom to top, comprising about 10 linear feet of nesting material — fine dried grasses and "old-man's beard" {Chionan- thus virginicus). The series of nests in- dicates usage of the tree cavity by one or more squirrels over a long period of time. Another nest was extracted from a hol- low tree northeast of Custer on 24 June 1967; approximately 538 cubic inches of
78
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
dried grasses and finely shredded bark constituted the nest material.
Foods of red squirrels in the Hills consist mainly of seeds of coniferous trees, but bracket fungi and acorns of bur oak also are consumed. A conifer cone is stripped of all its scales as the squirrels eat the ovules, dropping the gnawed core to the ground. These cut- tings generally accumulate beneath den or nest trees, forming large middens ( ac- cumulations of refuse about a dwelling place). Some middens that I have ex- amined were several feet in depth and exceeded 10 feet in diameter. Bailey ( 1888:436) noted that red squirrels abide in rocky bluffs and store cones in clefts among the rocks; he also reported ex- tracting tvi^o or three bushels of cones from a hollow pine that contained a nest. In addition to cone scales and cores, pine and spruce needles, gnawed twigs and bones, and other litter of food-remains comprise the bulk of such middens. A maze of underground burrows and blindly-ending pockets quite often pene- trate these deposits. In the northern Hills, cores of white spruce cones are more predominant in refuse piles than are those of ponderosa pine. Frequently, abandoned barns or feed-bins, and attics of deserted buildings are sites of vast middens and food caches.
Vernon Bailey obtained two lactating adult females southwest of Custer on 26 May 1894, and Ned Dearborn captured another at Redfern on 30 May 1910. Five of 16 females collected in June were nursing, and the uteri of three others evidenced 4 (3-5) placental scars; four additional adult females showed no signs of reproductive activity. Four pregnant red squirrels (the only ones encountered in this study) taken between 20 and 28 June carried 5 (4-6) embryos that were 21.5 (6-42) in crown-rump length. The average length of testes of 17 adult males captured in the period 1 to 15 June was 14.2 (8-22), whereas that of 18 males taken between 16 and 30 June was 17.3 (8-31). Fifteen of 26 females obtained in July were lactating and 11 were repro-
ductively inactive; the average testicular length of 18 July-taken adult males was 16.6 (5-25).
Utilizing the aging criteria of Layne (1954:251), 47 juvenile and subadult T. hudsonicus taken (between 10 June and 15 August) in the Hills were grouped into age classes. Time of capture was then projected backward to estimate time of birth of these individuals; 10.6 percent were born in late March, 29.8 percent in early April, 44.7 percent in late April, and 14.9 percent in early May. The occurrence of pregnant females in late June ( coincident with attainment of maximal summer testicular length in males ) intimates that some adult females in the Black Hills produce two litters, one in spring and the other in mid- summer, or else that some females breed later.
Layne (1954:232-235) described the seasonal change in pelage of Tamias- sciuriis hiidsoiiicus loquax in central New York; in general, my observations of molt in dakotensis, although somewhat less detailed, concur with his. Adults replace body pelage twice each year. Progress of molt is gradual and somewhat diffuse, usually lacking distinct lines. Adults col- lected in late May, throughout June, and in early July were in various stages of pelage replacement. Several lactating females still retained irregular patches of winter pelage on the rump in late June and early July. Two adult males cap- tured at Sundance on 18 August 1913 were in full summer pelage, as were six individuals obtained in October. An adult male obtained at Jewel Cave Na- tional Monument on 20 November 1967 was in process of autumnal molt; new hairs were manifest beneath the older pelage over the shoulders, sides and ven- ter. Layne ( loc. cit. ) noted that hairs on the tail of adults are replaced but once annually, during the fall molt.
Young red squirrels obtained in June generally were in fine soft juvenal pelage, whereas seven young captured in mid- July were in fresh subadult pelage ( simi- lar to adult summer pelage) and two
TURNER: MAMMALS OF THE BLACK HILLS
79
others were undergoing pelage replace- ment over most of the body.
A male taken 2 mi W Nemo on 2 Jul)" 1967 was parasitized externally by ticks, Dcnnacentor amlcrsoni Stil(\s, and another male from southwest of Sturgis harbored chiggers, Miy(itwni])icula eso- ensis (Susa and Ogota). In addition, red squirrels throughout the Black Hills were infc\sted with unidentified fleas ( un- fortunately, collected specimens of these ectoparasites were misplaced ) .
Comparison of 24 male and 31 female T. hudsonicus from the Black Hills re- vealed no secondary sexual dimorphism in coloration or external and cranial measurements. Eight squirrels (four males and four females) from north of
Sundance are somewliat paler (27.4± 2.68), and reflect reds (54.6±0.15) more intensely and greens (25.3±0.10) less intensely than typical lilack Hills repre- sentatives; conversely, seven individuals (three males and four females) from south of Tinton are comparatively dark (25.6±1.75), and reflect reds (51.7± 0.17) less intensely and greens (26.5±: 0.08) more intensely than do other Hills squirrels.
Tamiasciunis hudsonicus dakotensis differs from T. h. haileiji, which occurs west of the Black Hills, in being larger in most external and cranial dimensions, and more reddish in color ( reflecting red hues more intensely, Tables 12 and 13). It differs from T. h. fremonti, to the south-
Table 12. — Geographic variation in coloration of the mid-dorsal pelage of adult Tamiasciunis hudsonicus obtained in .summer from
the Northern Great Plains.
Number and sex
of specimens
averaged
55(245,31$)
Mean SD
6(25,4$)
Mean
SD
7 (35,45)
Mean
SD
27 (165,115)
Mean SD
4(25,25)
Mean SD
10(45,65) Mean SD
10(65,45)
Mean SD
d) u
G O
H o o
<a o
O 0)
CL,
OJ <u
a
|
<u |
(U |
|
o |
t> |
|
c |
|
|
a o |
<D O |
|
<v <u |
o a; OJ |
|
OJ « |
|
|
So^ |
T. h. dakotensis. Black Hills, South Dakota and Wyoming
26.0 53.3 25.8 20.9 ±2.19 ±0.16 ±0.10 ±0.13
T. h. dakotensis, Converse County, Wyoming
30.7 53.9 25.3 20.8
±2.77 ±0.18 ±0.13 ±0.08
T. /). dakotensis. Carter County, Montana
20.6 51.5 26.7 2L8
±3.36 ±0.36 ±0.19 ±0.22
T. h. bailetji, Johnson and Big Horn
counties, Wyoming
22.3 50.1 27.2 22.7
±1.92 ±0.15 ±0.09 ±0.12
T. h. hailetji, southeastern Albany County,
Wyoming
27.6 49.3 27.2 23.5
±3.35 ±0.14 ±0.07 ±0.12
T. h. bailetji, Fergus County, Montana
25.1 49.4 26.9 23.7 ±2.32 ±0.19 ±0.13 ±0.14
T. h. fremonti. Carbon and southwestern
Albany counties, Wyoming
23.0 48.3 27.8 23.9
±1.45 ±0.14 ±0.14 ±0.97
so
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Table 13. — Geographic variation in selected external and cranial measurements of Tamiasciunis hudsonicus from the Northern Great Plains.
adult
|
Number and sex |
c |
i-C |
|
of specimens |
iS M |
4~* |
|
averaged |
o c |
bc
o c
o
XI
||
G
O
o -^
S rt O
55 (24 5,31$ )
Mean
SD
6 (25,4$)
Mean SD
7 (3,5,45)
Mean SD
27 (16 5,115)
Mean
SD
4 (25,25 )
Mean
SD
10 (45,65) Mean SD
10(65,45) Mean SD
T. h. dakotensis. Black Hills, South Dakota and Wyoming
347.7 27.4 291.9 50.3 19.0 29.1 14.3 ±18.40 ±2.34 ±27.70 ±0.89 ±0.52 ±0.83 ±0.44
T. h. dakotensis. Converse County, Wyoming 343.5 26.0 250.3 48.6 18.8 28.4 14.5
±17.24 ±0.89 ±15.71 ±0.73 ±0.56 ±0.26 ±0.64
T. h. dakotensis, Carter County, Montana 340.9 27.4 289.4 49.9 19.3 29.0 14.3
±13.91 ±1.15 ±18.09 ±1.23 ±0.49 ±0.86 ±0.35
T. li. haileiji, Johnson and Big Horn counties, Wyoming .328.9 25.8 245.4 49.0 18.4 28.1 14.6
±20.92 ±1.80 ±39.24 ±0.93 ±0.53 ±0.93 ±0.42
T. h. haiJetji, southeastern Albany County, Wyoming 338.0 23.7 238.5 49.3 19.3 29.2 15.2
±14.44 ±5.91 ±36.01 ±0.40 ±0.50 ±0.43 ±0.63
T. h. baileyi, Fergus County, Montana
342.8 25.4 284.3 49.0 18.5 28.6 14.5 ±8.01 ±2.46 ±44.34 ±1.50 ±0.59 ±0.73 ±0.21
9.0 ±0.31
8.0 ±0.37
9.0 ±0.27
8.7 ±0.29
8.5
±0.13
8.6 :0.20
T. h. ficmonti. Carbon and southwestern Albany counties, Wyoming
327.8 26.2 240.5 48.7 19.4 28.5 15.1 8.4
±12.72 ±2.39 ±18.06 ±0.57 ±0.47 ±0.67 ±0.53 ±0.24
west, in the same manner bnt to a greater extent. Specimens of fremonti reflect greens and blues much more intensely than do those of dakotensis. When tone and hue are analyzed separately (see methods and materials ) , red squirrels are found to vary considerably in tone of color, but the three races discussed here do not differ significantly in this param- eter; rather, the races are differentiated on basis of reflection of the various hues.
Because subspecific identification of red squirrels from several areas in south- eastern Montana and Wyoming is open to question (Hoffmann and Jones, 1970: 374, Fig. 7), these outlying populations also were examined (Tables 12 and 13). Specimens taken 21.5 mi S, 24.5 mi W Douglas, Converse Co., Wyoming, repre- sent intergrades between bailey i and dakotensis, but on basis of color are
assigned here to the latter. Individuals from a little farther south (southeastern Albany County), however, are clearly haileiji. Red squirrels from Carter County, Montana, also are here desig- nated as dakotensis on the basis of size and color.
Layne (1954:230) observed the fre- quency in occurrence of the minute first upper premolar of T. h. loqiiax to be 67.7 percent in males and 70.3 percent in fe- males, and indicated that Bangs (1896) noted this tiny tooth was absent in more than 50 percent of T. J}, liudsonicus ex- amined. (In consulting the literature, I find no such statement by Bangs.) A peg-like first premolar was visible in the skulls of IS percent of 55 adult males and in 25 percent of 40 adult females of dakotensis examined from the Black Hills. Measurements additional to those
TURNER: MAMMALS OF THE BLACK HILLS
81
in Table 13 of 55 Hills specimens are: tail length, 137.6 ±9.44; liind foot length, 52.0±4.00; and skull depth, 21.8+0.80.
Specimens examined (295).— SOUTH DA- KOTA: Lawrence County: Spearfish, 6 (5 SDSU, 1 USNM); Big Spearfish Canyon, 3 mi S Spearfish, 4200 ft, 1; Big Spearfish Canyon,
5 mi S, 2 mi E Spearfish, 4800 ft, 1; White- wood, 4 (USD); Tinton, 2 mi N, 13 mi W Lead, 6000 ft, 4; 2 mi S Tinton, 6100 ft, 22; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 6; Little Spearfish Canyon, Savoy, 1 (USNxM); Annie Creek, 6500 ft, 1 (USNM);
8 mi W Lead, 2 (SDSU); Deadwood, 6 (USNM); 4 mi SW Lead, 1 (NRW); Terry Peak, 1 (NRW); 1 mi S, 7 mi W Lake Rou- baix, 1; 4 mi N, 1 mi W Nemo, 4900 ft, 1; 2 mi W Nemo, 4700 ft, 10; 1 mi E Nemo, 4700 ft, 1. Meade County: Vanocker Canyon, 4 mi S, 1 mi W Stmgis, 4600 ft, 1; Haven Dams,
6 mi S, 1.5 mi W Sturgis, 4600 ft, 8; 9 mi S,
1 mi W Sturgis, 4800 ft, 3. Pennington County: Rochford, 6 (UMMZ); 6 mi N, 2 mi W Silver Cit>', 1; Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 17; Beaver Creek, 3 mi N,
9 mi W Deerfield, 6500 ft, 2; 5 mi S, 1 mi W Pactola Lake, 1 (SDSU); Pactola Dam, 1 (SDSU); 10 mi W Rapid City, 1 (SDSU); South Canyon, 4 mi W Rapid City, 6 ( AMNH ) ; Rapid City, 1 (SDSU); Dark Canyon, 2 mi S, 4 mi W Rapid City, 3 (AMNH); Deerfield Lake, 6400 ft, 2; Deerfield, 1 (SDSU); Red- fern, 2 (USNM); 5 mi S, 1 mi W Pactola Lake, 1 (SDSU); 1 mi W Rockerville, 1 (NRW); Spring Creek, south of Rapid City,
2 (MHM); Ditch Creek, 14 mi VJ Hill City, 6400 ft, 37; Glendale, 8 (7 AMNH, 1 FMNH); Hill City, 3 (USNM); 2 mi SW Medicine Mountain, 1 (SDSU); 3-4 mi SE Hill City, 5300-5400 ft, 16 (UMMZ); Summit Peak, 5500 ft, 1 (UMMZ); Horsethief Lake, 3 mi W Mount Rushmore, 5100 ft, 1; near Mount Rushmore, 1 (UMMZ); 1 mi S Keystone, 1 (NRW); 20 mi N Elk Moimtain, 6000 ft, 1 (USNM); 17 mi NW Custer, 2 (UMMZ); 16 mi NW Custer, 3 (UMMZ); unspecified locafity, 4 (1 SDSU, 3 UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, .5.300 ft, 2 (FMNH); 3.5 mi S, 0.5 mi W Key- stone, 5000 ft, 7; 5.75 mi N, 5.75 mi E Custer, 5220 ft, 8; Roby Springs, 4 mi N, 22 mi W Custer, 5400 ft, 6; 4 ini N, 14 mi W Custer, 2; 4 mi N, 4.5 mi E Custer, 5400 ft, 1; Tepee Canyon, 14 mi W Custer, 3 (USNM); Custer,
7 (1 SDSU, 6 USNM); Harvey National For- est, 3 mi E Custer, 2 (UMMZ); Custer State Park, 5 (UMMZ); Squaw [=Grace Coolidge] Creek, 5 (3 AMNH, 2 FMNH); Jewel Cave National Monument, 2.5 mi S, 12 mi W Custer, 5400 ft, 1; 7 mi SW Custer, 1 (SDSU); Elk Mountain, 1 (USNM), unspecified locafity, 1 (UMMZ). Fall River County: 0.5 mi S, 1.5
mi W Minnekahta, 4200 ft, 1. Unsj}ecified county: Black Hills, 3 (1 NRW, 1 SDSU, 1
USNM).
WYOMING: Crook County: Devils Tower, 4 (USNM); 15 mi N Sundance, 5500 ft, 1; 15 mi ENE Sundance, 3825 ft, 7; 3 mi NW Sundance, 5900 ft, 16; Sundance, 2 (USNM); unspecified locality, 1 (SDSU).
Additional records.— SOUTH DAKOTA: Lawrence County: Lead (Chambers, 1948:8). Custer County: 16 mi SW Custer (USBS files). WYOMING: Weston County: head Cheyenne River (J. A. Allen, 1877:689).
Glaucomys sabrinus bangsi (Rhoads)
Northern Flying Squirrel
Scittropterus alpinus bangsi Rlioads, 1897, Proc. Acad. Nat. Sci. Philadelphia, 49:321, 19 July (type locality, Raymond, Bear Lake Co., Idaho).
Glaucomys sabrinus bangsi — A. H. Howell, 1918, N. Amer. Fauna, 44:38, 13 June.
Glaucomys sabrinus occurs in mon- tane habitats of the western United States, and in boreal forest areas of north- ern North America. In the Black Hills, this squirrel is most abundant in the spruce forests of moist canyon systems on the boreal cap, but ranges down to about 4500 feet. Specimens are lacking for Meade, Custer, and Fall River coun- ties; however, residents report that this species is a common mammal through- out the study area, and is active the year around. Grinnell (1875:81) first noted flying squirrels in the Hills, under the name Pteromys alpinus.
Northern flying squirrels are some- what gregarious in the colder months. J. A. King captured 10 individuals in a spruce-filled canyon southeast of Hill City between 11 December and 29 March; three of these were obtained from the same tree. These specimens were taken in rat traps that were at- tached to trees about seven feet above ground level and baited with peanuts. South of Tinton, remains of a flying squirrel were found in the center of a midden of pine cones at the base of a dead tree.
Most of the recently collected speci- mens have been obtained by tapping on dead hollow trees and shooting the squir-
82
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
rels as they exited from the den. Trees selected as nest sites usually are high on slopes wooded in pine and scattered spruce and aspen. The average height of five nest trees was 18.6 (15 to 25) feet, and all were about one and a half feet in diameter; openings to these nests were 12.8 (10 to 18) feet above ground surface. The oval-shaped entrance to one nest was 57 by 67 millimeters in diameter; depth and width of the nest cavity were 145 and 100 millimeters, re- spectively; the spherical nest was com- posed of fine grass and "old-man's beard." Flying squirrels were obtained from nests east of Buckliorn, Wyoming, on 18 July 1951 (one lactating adult female and four juveniles, of which two were males and two were females), and west of Nemo, South Dakota, on 29 June 1967 (one lactating adult female and two juveniles, one male and one female), on
4 July 1967 (one adult male), and on
5 July 1967 (one lactating adult female and three juveniles, of which two were males and one was a female).
Reproductive data essentially are lacking for this species in the Black Hills except for the nest-taken individ- uals mentioned above. Testes of an adult male taken on 4 July were 11 in length. Young flying squirrels are in evidence through August. These have a fine short Juvenal pelage that generally is darker than that of adults; the tail is blackish rather than cinnamon. Glaucomys sa- Ijrinus from the Black Hills were para- sitized externally by an unidentified flea.
A. H. Howell (1918:38) referred two specimens of G. sahrinus from the Bear Lodge Mountains to the subspecies canescens, indicating that these individ- uals approached bangsi in skull charac- ters and in a slightly more vinaceous coloring on the upperparts. When addi- tional specimens were made available through his own field work. King (1951: 469) noted that representatives of Glau- comys from the Black Hills differed sig- nificantly from canescens of North Da- kota, but closely resembled three speci- mens of bangsi from Park and Fremont
counties, in Wyoming; Long (1965:599) later concurred with King's assignment of flying squirrels in the Black Hills to G. s. bangsi.
Neither secondary sexual character- istics nor age groups of adults (age was determined in the manner of King, loc. cit.) evidenced significant variation. In- dividuals from tlie Hills were compared with specimens from Park, Teton, and Sublette counties, in western Wyoming. Mid-dorsal pelage of animals from the Black Hills is slightly darker in tone, and reflects reds more intensely and greens less intensely than those from western Wyoming; however, tliese differences are nonsignificant. Means and standard de- viations of these color parameters are given for six adults (two males and four females) from the Hills, followed by those for nine adults ( four males and five females) from the western Wyoming: color tone, 39.1±2.94, 39.5±4.78; per- cent red reflectance, 51.2±0.12, 50.0± 0.28; percent green reflectance, 26. 1± 0.07, 27.3±0.16; percent blue reflectance, 22.7±0.08, 22.7±0.16.
External and cranial dimensions of representatives from the Hills examined in the present study agree closely with those of 12 adults measured by King (loc. cit.). However, specimens from the study area are significantly smaller than those from western Wyoming in total length, tail length, ear length, great- est length of skull, and zygomatic breadth. Means and standard deviations of sLx adults (two males and four fe- males) from the Hills, followed by those of nine adults (four males and five fe- males) from western Wyoming, are: total length, 300.8±11.43, 329.4±9.69; tail length, 133.7±5.28, 150.0±4.08; hind foot length, 40.0±3.35, 41.3±1.11; ear length, 23.3±4.27, 27.3±1.11; weight, 149.8±23.48, 167.5+33.78; great- est length of skiUl, 40.4±0.69, 41.6±0.81; zygomatic breadth, 24.3 ±0.25, 25.2± 0^50; braincase breadth, 19.3±0.33, 19.5 ±0.22; postorbital breadth, 9.0±0.39, 9.0±0.50; interorbital breadth, 7.7±
TURNER: MAMMALS OF THE BLACK HILLS
83
0.30, 8.0 ±0.63; rostral longtli, 13.5 ± 0.32, 13.9 ±0.59; length of maxillary tooth-row, 8.1 ±0.14, 8.1 ±0.35; cranial depth, 17.6 ±0.51, 18.1 ±0.55; ratio of cranial depth to breadth of braincase, 0.92 (range: 0.91-0.95), 0.93 (range: 0.89-0.96).
The smaller size, slightly different coloration, and isolation of G. sahrinus in the Black Hills ( about 245 miles west of populations in Park County, Wyom- ing ) warrant further investigation before an accurate assignment to a subspecies can be made. For the present time, 1 tentatively retain the subspecific name ])an(isi for the flying squirrels in the Black Hills.
Specimens examined (31). — SOUTH DA- KOTA: Lawrence County: Little Spearfish Canyon, Timon Campground, 1 (USNM); 2 mi S Tinton, 6100 ft, 1; 2 mi W Nemo, 4700 ft, 7. Pennington County: 1 mi N Sheridan Lake, 1 (USNM); Castle Creek, 6500 ft, 2 (UMMZ); 4 mi SE Hill City, 53-5400 ft, 16 (UMMZ); T. 1 N, R. 6 E, sec. 8, 1; 17 mi NW Custer, 1 (UMMZ); 16 mi NW Custer,
2 (UMMZ).
WYOMING: Weston County: 0.5 mi N,
3 mi E Buckhorn, 6200 ft, 5.
Additional records. — WYOMING: Crook County: middle fork Hay Creek, Bear Lodge Mountains (A. H. Howell, 1918:38).
Family GEOMYIDAE— Pocket Gophers
This family of fossorial rodents has undergone its entire evolutionary history in North and Central America. Pocket gophers, so named because of the fur- lined cheek pouches that open exter- nally on either side of the mouth, are rep- resented by but one species in the Black Hills; a second species {Geomijs hursar- ius) has been reported erroneously from the area. Food habits of pocket gophers and their extensive subterranean burrow systems, usually marked by mounds of earth near each entrance, present some economic problems to agricultural and livestock interests; thus control methods such as extensive trapping or poisoning programs have been carried out by ranchers in the Hills region.
Thomomys talpoides nebulosus V. Bailey
Northern Pocket Gopher
Thomomys talpoides nebulosus V. Bailey, 1914, Proc. Biol. Soc. Washington, 27:116, 10 July (type locality. Jack Boyden's Ranch, Sand Creek Canyon, 15 mi NE Sundance, 3750 ft, Crook Co., Wyoming).
The northern pocket gopher is wide- spread in the Black Hills and common where local environmental factors are favorable. The species inhabits montane meadows and parklands, disturbed areas of secondary vegetative growth, and croplands, but is most abundant in heav- ily grazed valley pastureland. Specimens are available from throughout the Hills save from Fall River County, but the earthen mounds characteristic of gophers evidence their presence there. Based on specimens at hand, the altitudinal range of T. t. nebulosus, a subspecies endemic to the Black Hills, extends from 3500 to 6500 feet. Among 146 individuals ex- amined of which the sex was known, oiily 41 are males, suggesting that females may be more readily captured in traps. Thomomys talpoides was first reported from the Hills by Bailey (1888:450).
The holotype, four allotypes, and specimens from Sundance were taken by Vernon Bailey late in August 1913 in potato fields. Those obtained northeast of Beulah, Wyoming, early in July 1947 were captured in fields of alfalfa. Speci- mens from Palmer Gulch were trapped in winter in a valley hayfield, between a pine forest and a small stream. Four females were taken in August 1934 in a montane meadow on Doran's Ranch, 3 mi E Custer by Stebler ( 1939:390) where prominent plants were the short grasses, but clumps of sagebrush, goldenrod, but- ter and eggs, and asters were conspicu- ous.
Fifteen individuals from Nemo (late June and early July 1967) were trapped in a grazed pasture of bluegrass, clover, and scattered forbs. Three of eight go- phers trapped east of Four Corners early in July 1965 were taken in a burned area on a high ridge on which secondary re- generation of ponderosa pine and many
84
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
kinds of forbs had occurred; the others were captured in a pasture of bluegrass and clover. No specimens have been ob- tained in heavily forested areas.
In early July 1965, a member of a field party from the Museum of Natural History, H. H. Genoways, studied burrow systems of T. t. nebulosus and their dis- tribution in Beaver Creek Valley, Penn- ington County. Mounds were located in dense, sodlike grasses where the soil was extremely dark, moist, and fairly free of rocks, as well as in areas of dry rocky soils that contained much organic material. Burrows generally were lo- cated at the edge of mounds. The fol- lowing average measurements (in milli- meters) are of 16 tunnel systems in valley soils: depth from opening of mound to bottom of tunnel, 170 ( 110- 250); width of tunnel, 65 (50-80); and height of mound above ground level, 75 (60-90). Burrows located in rocky soils generally were much shallower, and sys- tems excavated by subadults were at most 80 millimeters below ground level and were situated in relatively deep black soils.
This species often is active in the early afternoon. A female was obtained in Weston County on 7 July 1965 at 15:00 hrs (MDT), shortly after traps were set, and I captured two juveniles about 16:00 hrs (MDT) in snap-traps that were set in a montane meadow southwest of Spearfish on 7 August 1967. Although most of the specimens examined were taken in the summer months, J. A. King obtained individuals from November 1945 through April 1946 in Pennington County. Mounds and tunnel systems formed under the snow usually are evi- dent in spring, a further indication that the northern pocket gopher is active the year around in the Hills.
Testicular length of an adult male captured on 4 July 1967 was 12. Only two pregnant females were observed, one taken on 3 July 1967 at Nemo, South Dakota, and the other collected on 4 July 1965, near Four Corners, Wyoming; each carried four embryos that were 9
and 12 in crown-rump length, respec- tively. Enlarged uteri, possibly indicat- ing recent implantation were noted in three females obtained in early July 1965 in Beaver Creek Valley; two others were lactating. Placental scars (7, 3, and 3) were evident in three females taken in Lawrence County in 1967 on 29 June, 3 July, and 29 July, respectively. Females probably breed once annually ( from mid- March through mid- June) and produce three or four young; thus, the occurrence of seven placental scars in late June is of interest. Placental scars in pocket gophers generally remain two months after parturition (Tryon, 1947:18) but the number of live-born young may be smaller than the embryo count; addi- tional scars probably indicate resorption of some embryos rather than production of two litters at this early date. Hansen (1960:331) reported resorption of about four percent of all embryos examined in a study of T. talpoides in Colorado.
Young of the year were captured first in mid-June in the Black Hills and are fairly common in collections made in July; for example, 10 of 15 individuals obtained at Nemo from 29 June to 4 July 1967 are juveniles, as are five of six specimens taken at Beulah on 2 July 1947. Subadults attain about three- quarters of adult size by late July and early August, although younger individ- uals are still taken occasionally. Two traps placed in the same burrow system at Nemo took an adult female and a juvenile female on 3 July 1967.
Bailey (1915:18) described the proc- ess of molt in T. talpoides. Progress of the crescentic shaped molt wave is from the rostrum, toward the rump. Young taken in late June and early July were in a fine, juvcnal pelage; many subadults trapped late in July were in post-juvenal molt. Two simultaneous molt lines were of common occurrence on spring-taken adult specimens, and five individuals evi- dence three successive molt lines in progress. Long winter pelage remain on the rump of most specimens until mid- July. A female taken by Merritt Cary
TURNER: MAMMALS OF THE BLACK HILLS
85
on Elk Mountain, on 19 November 1901 was undergoing molt to winter pelage, whereas three others obtained there about this time were in fresh winter pelage. Four specimens from tlie iilaek Hills ha\e patches of wliite hair on the shoulder or dorsum, and a female from Nemo has a white star-shaped area on the forehead.
A female from Pennington County (2 July 1965) was parasitized externally by two kinds of lice, Geomyduecus tho- monujus (McGregor), and a new species currently being described by R. D. Price and associates, University of Minnesota.
Analysis of nongeographic variation in northern pocket gophers in the Black Hills revealed no secondary sexual di- morphism in external or cranial dimen- sions, but tone of color of seven adult males collected in the summer (2S.6± 4.38) is significantly paler than that of 20 adult females (23.9±4.30). Because of the topographic diversity in the moun- tainous terrain, and due to the relatively sedentary nature of pocket gophers, lo- calized populations display great vari- ability in the Hills. For example, indi- viduals from Nemo are darker in tone, reflect blues more intensely, and have greater lengths of skull, rostrum, and
maxillary tooth-row than other popula- tions in the Hills.
TJioniomys talpoides nebulosus in the Black Hills resembles T. t. InilJattis of the plains to the west, north and north- east, but is significantly darker in tone and larger in most external and cranial dimensions (see Table 14). Specimens from outlying areas of the Hills (Bear Lodge Mountains, Devils Tower, Four Comers, \'icinity of Beulah and near Buckliorn, in Wyoming, and Rapid City, in South Dakota) are paler than typical nebulosus, showing a tendency toward huUatus, but are best assigned to the former. Thomoimjs talpoides nebulosus differs from T. t. pierreicolus of the plains to the east and south in the same manner as described above for T. t. bullatus, but to a greater extent. Selected measure- ments of seven adult males and 24 adult females from the Hills are: percent red reflectance, 45.9 ±0.24; percent green re- flectance, 27.6±0.13; percent blue reflect- ance, 26.5±0.20; body length, 226.3± 9.34; tail length, 66.6±6.58; hind foot length, 29.1 ±2.36; ear length, 6.8± 0.88; weight, 140.6 ±19.78; greatest length of skull, 38.9±1.61; zygomatic breadth, 23.2 ±0.94; rostral length, 17.1 ±1.09; interorbital constriction, 6.6±
Table 14. — Geographic variation in selected color, and external and cranial measurements of adult
Thomoimjs talpoides from the Northern Great Plains.
|
Number and sex of specimens averaged |
Tone of color |
Total body length |
Tail length Hind foot length Weight Greatest length of skull Rostral length |
Length of maxillary tooth-row |
Skull depth |
|
31 (75,245) Mean SD |
25.1 ±4.72 |
T.i 266.3 ±9.34 |
'. nebulosus. Black Hills, South Dakota and Wyoming 66.6 29.1 140.6 38.9 17.1 7.9 ±6.58 ±2.36 ±19.78 ±1.61 ±1.09 ±0.39 |
15.3 ±0.80 |
|
|
13(6<5,79 ) Mean SD |
37.0 ±6.85 |
213.7 ±10.99 |
T. t. bullatus, Harding County, South Dakota 64.5 27.5 130.5 38.0 16.2 ±6.13 ±1.39 ±22.28 ±1.85 ±0.90 |
7.6 ±0.32 |
14.5 ±0.49 |
|
9(15,8?) Mean SD |
33.2 ±1.37 |
212.9 ±9.20 |
T. t. bullatus, Campbell County, Wyoming 60.9 27.9 109.2 37.4 16.2 ±4.23 ±1.27 ±11.34 ±0.86 ±0.58 |
7.7 ±0.32 |
14.7 ±0.59 |
|
4(15,39) Mean SD |
32.1 ±3.82 |
199.7 ±4.27 |
T. t. pierreicolus, Dawes County, Nebraska 54.2 27.0 92.8 35.5 14.4 ±2.63 ±1.15 ±5.92 ±0.96 ±0.54 |
7.1 ±0.26 |
14.3 ±0.46 |
86
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
0.34; length of maxillary tooth-rr)w, 7.9 ±0.39; depth of skull, 15.3+O.SO.
Specimens examined (158). — SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- yon, 9 mi S, 3 mi W Spearfish, 5000 ft, 2; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 2; 2 mi S Tinton, 6100 ft, 2; 2 mi N, 4 mi W Nemo, 5100 ft, 3; Nemo, 5700 ft, 15; un- specified locality, 1 (NRW). Meade County: Fort Meade, 1 (USNM). Pennington County: Rapid City, 2 (USNM); Spring Creek, south of Rapid City, 2 (MHM); Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 3; Beaver Creek, 3 mi N, 9 mi W Deerfield, 6500 ft, 3; 20 mi N Elk Mountain, 2 (USNM); Ditch Creek Valley, 14 mi W Hill City, 6400 ft, 1;
2 mi S, 13 mi W Hill City, 1; Palmer Gulch,
3 mi SE Hill Cit>', 5300 ft, 14 (UMMZ); Tiger- ville, near Hill City, 2 (USNM); Redfern, 4 (USNM); 16 mi NW Custer, 3 (UMMZ); Harney National Forest, 3 mi E Custer, 4 (UMMZ); unspecified locality, 6 (UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, 5300 ft, 4 (FMNH); 3.5 mi S, 0.5 mi W Keystone, 5000 ft, 1; Bull Springs, 2 mi N, 9 mi W Custer, 6 (UMMZ); Custer, 5 (4 USNM, 1 AMNH); Mayo, 3 (USD); Camp- bell's Ranch, Elk Mountain, 4800 ft, 1 (USNM); Elk Mountain, 7 (USNM); 1 mi N Wind Cave, Wind Cave National Park, 1 (WCNP); Wind Cave National Park, 1 (USNM); Beaver Creek, 1 (USNM).
WYOMING: Crook County: 1 mi E, 0.5 mi N Beulah, 3550 ft, 14; Beulah, 3500 ft, 6; Bear Lodge Mountains, 6200 ft, 6 (USNM); Devils Tower, 1 (USNM); 1.5 mi NW Sundance, 5000 ft, 4; Sundance, 2 (USNM); Rattlesnake Creek, 6000 ft, 2 (USNM); Sand Creek, 5 mi above mouth of canyon, 5 (USNM). Weston County: 1.5 mi E Buckhorn, 61.50 ft, 2; 4 mi E Four Corners, 8; Newcastle, 5 (USNM).
Additional record.— SOUTH DAKOTA: Lawrence County: Deadwood (Bailey, 1888: 450).
Family HETEROMYIDAE— Pocket Mice axd Kaxgaroo Rats
Three species ( two genera ) of hetero- myids are represented in the mammalian fauna of the Black Hills; a fourth species occurs nearby and its status in the Hills region remains uncertain. All members of this family possess external, fur-lined cheek pouches, and to varying degrees are nocturnal, fossorial, and saltatorial. These rodents inhabit the interface of the foothills and surrounding semiarid plains, particularly on fine loamy soils of the Sand Hill Regosol Soil Subassocia-
tion along the South Dakota-Wyoming border.
Perognathus fasciatus olivaceogriseus
Swenk
Olive-backed Pocket Mouse
Perognathus flavescens olivaceogriseus Swenk, 1940, Missouri Valley Fauna, 3:6, 5 June (type locality. Little Bordeaux Creek, sec. 14, T. 33 N, R. 48 W, 3 mi E Chadron, Dawes Co., Nebraska).
Perognathus fasciatus olivaceogriseus — Jones, 1953, Univ. Kansas Publ., Mus. Nat. Hist., 5:520, 1 August.
The olive-backed pocket mouse in- Iiabits gravelly soils on the high plains peripheral to the Black Hills, and oc- casionally, individuals are taken in the foothill transition zone. The species has been captured only at four localities within the Hills of which two are in South Dakota, and two are in Wyoming. Altitudinal range as indicated by these specimens is up to 4800 feet.
Most available specimens of this spe- cies were obtained in the Hills area in the early part of the present century, but thev first were reported from the area by Jones (1953:522): Merritt Caiy captured five females in early October 1903 in Custer County; Vernon Bailey obtained a male near Newcastle, on the western border of the Hills, on 24 May 1894, and in August 1913 found a mum- mified individual of unknown sex near Sundance, Wyoming.
I obtained two specimens while trap- ping in Wind Cave National Park in 1968; a female was taken on 29 July (partially eaten in the trap by a chip- munk) and a male on 28 August. Both indi\'iduals were captured in snap-traps baited with rolled oats and placed in sparse grass at the junction of a fence row and a gravel road; the fence en- circled a sewage-settling pond at the head of Wind Cave Canyon. Micwtus ochro<i,aster, Peromyscus maniculatiis, Reithroclontomys meg,aJotis, Perognathus Jiispidus, and Eutamias minimus were taken in the same trapline.
TURNER: MAMMALS OF THE BLACK HILLS
87
Tlie female taken in 196(S was neither pregnant nor laetating; reprodnctive data are unavailable for those females cap- tined by Gary. Testes of the adult male from Wind Ca\e Canyon were 3 in length.
Specimens examined (8). — SOUTH DA- KOTA: Custer County: Campbell's Ranch, Elk Mountain, 4800 ft, 5 (USNM); Wind Cave Canyon, Wind Cave National Park, 4100 ft, 1.
WTOMLNG: Crook County: Sundance, 1 (USNM). Weston County: Newcastle, 1 (USNM).
Perognathus hispidus paradoxus
Merriam
Hispid Pocket Mouse
Ferognathus paradoxus Merriam, 1889, N. Amer. Fauna, 1:24, 25 October (type lo- cality. Banner, Trego Co., Kansas).
Ferognathus hispidus paradoxus — Osgood, 1900, N. Amer. Fauna, 18:44, 20 September.
The hispid pocket mouse ranges across the short and mixed grass prairie of western South Dakota and eastern Wyoming (Glass, 1947:179). Although imcommon, it has been taken in the foot- hill transition zone of the Black Hills at altitudes up to 4800 feet. Specimens ha\'e been obtained only in Guster and Fall River counties.
Three females were captured by Mer- ritt Gary on Gampbell's Ranch, Guster Gounty, in the early 1900's, and a field part}' from The University of Kansas found a male dead along a road 3 mi E Pringle in mid-June 1961. A juvenal fe- male was trapped southwest of Minne- kahta in mid-June 1967, along an over- grown fence row that was bordered on both sides by barren agricultural fields. The fence was paralleled by a strip of dense vegetation about 10 feet in width, consisting of Japanese brome, western wheat grass, bluegrass, and a few forbs.
Two males were taken in late sum- mer of 1937 in Wind Gave National Park by E. Suter. I obtained a female in the Park in mid-August 1967 and a male at the head of Wind Gave Ganyon in late August 1968. The female was carr\dng unidentified seeds in her cheek pouches
and was trapped along a high fence that separates the National Park from Guster State Park. Vegetation, mainly grasses, on the Wind Gave side of the fence was tall and dense. Due to overgrazing by bison in Guster State Park, vegetation was sparse and short on that side of the fence. The male was trapped in an open habitat of sparse vegetation, where the soil was rocky, being similar to a gravel pavement. Microtus ochro<i,aster, Pero- mysctts maniculatiis, Reithrodontomys 7negalotis, Perognathus fasciatus, and Eu- tamias minimus were taken in associa- tion with P. h. paradoxus.
Reproductive data concerning fe- males captured by Gary are unavailable; however, the female collected on 18 Au- gust 1967 carried six embryos that each were 15 in crown-rump length, and testes of the adult male taken on 27 August 1968 were 5 in length. The fuvenal female ( 17 June 1967 ) was in fresh pelage that was fine in comparison to the rather coarse fur of adults. The male from Wind Gave Ganyon was parasitized ex- ternally by ticks, Ixodes spinipajpis Had- wen and Nuttall.
Specimens examined (9). — SOUTH DA- KOTA: Custer County: Campbell's Ranch, Elk Mountain, 4800 ft, 3 (USNM); 3 mi E Pringle, 1; Wind Cave National Park, 2 (WCNP); 6 mi N, 1 mi E Wind Cave, Wind Cave National Park, 4400 ft, 1; Wind Cave Canyon, Wind Cave National Park, 4100 ft, 1. Fall River County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 1.
Dipodomys ordii luteolus (Goldman) Ord's Kangaroo Rat
Ferodipus ordii luteolus Goldman, 1917, Proc. Biol. Soc. Washington, 30:112, 23 May (type locality, Casper, Natrona Co., Wyo- ming).
Dipodomys ordii luteolus — Grinnell, 1921, Jour. Mamm., 2:96, 2 May.
Ord's kangaroo rat occurs on the plains that surround the Black Hills, and on sandy soils with sparse ground cover and an abundance of seed plants. Oc- casionally, these rats also range into the foothill transition zone. Merritt Gary collected a male and a female on Gamp- bell's Ranch at Elk Mountain early in
88
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
October 1903 and N. R. Whitney ob- tained a male south of Cascade Springs, Fall River County, on 8 February 1957. Two other individuals were taken along the eastern periphery of the Hills, south of Rapid City, sometime between 1899 and 1911 by Henry Behrens. D. G. Adolphson recently obtained skulls of this species from barn owl pellets col- lected 15 mi S Hot Springs ( Martin, 1971a: 4; 1971b).
The female from Elk Mountain, taken on 10 October, evidenced molt on the dorsum; no other life history data are at hand for the specimens examined.
Specitnem examined (5). — SOUTH DA- KOTA: Penniufiton County: Spring Creek, south of Rapid City, 2 (MHM). Custer Coiintt/: Campbell's Ranch, Elk Mountain, 4800 ft, 2 (USNM). Fall River County: south of Cascade Springs, 1 (NRW).
Dipodomys ordii terrosus Hoffmeister Ord's Kangaroo Rat
Dipodomys ordii terrosus Hoffmeister, 1942, Proc. Biol Soc. Washington, 55:165, 31 De- cember ( type locality, Yellowstone River, 5 mi W Forsyth, 2750 ft, Rosebud Co., Montana ) .
Habits of this heteromyid are similar to those of the preceding subspecies. Vernon Bailey obtained three males and four females near Newcastle, Wyoming in late May 1894. When he examined kangaroo rats from the Northern Great Plains region, Setzer (1949:524 and 543) designated those specimens from Elk Mountain (in the Black Hills proper) as D. o. hiteohis and indicated that other individuals from near Newcastle (on the western periphery of the Hills ) were in- tergrades between D. o. hiteohis and D. o. terrosus, but were assignable to the latter. D. o. terrosus is discernibly darker and larger than hiteohis except for a shorter tail and external ear pinna (Long, 1965:618).
Further collecting is requisite along the western border of the Hills to pro- vide critical data concerning the status of these two subspecies in that region. Until further specimens are available, I follow Setzer.
Specimens examined ( 7 ) . — WYOM ING : Weston County: Newcastle, 7 (USNM).
Family CASTORIDAE— Beavers
The beaver is the largest rodent in North America. Prior to the appearance of European man in the region, beaver were of great abundance and played a major role in luring early adventurers to the Black Hills. For example, expeditions of the Verendrye brothers in 1743, the American Fur Company in 1811, Jedc- diah Smith in 1823, and untold number of traders, trappers, and mountain men in the early 1800's mainly were in search of beaver and the premium prices paid for beaver pelts at that time.
Morgan (1868:137) indicated that beavers migrate each year, particularly when an area becomes overstocked. He noted that an annual migration down the Missouri River usually occurred in June, and wrote as follows: "A trapper whom I met on the Missouri River in 1862, below Fort Pierre, in Nebraska [present day South Dakota], informed me that beavers were then (May 27) coming down the river; that he saw them dailv and had taken over fifty." Inci- dentally, Morgan (1868:209) gave a de- scription of the natural vegetation along the Missouri River as it appeared in the 1860's, before modification by white man.
As evidenced by numerous houses and dams along most streams, beaver were abundant in the Black Hills at the time of the Custer Expedition in 1874 (Grinnell, 1875:82; Ludlow, 1875:15; O'Hara, 1929:252) and the Newton- Jenney Expedition in 1875 (Dodge, 1876: 123). Increased colonization caused re- ductions in numbers of beavers, and Bailey (1888:441) reported that in the autumn of 1887, only a few were "still found in places far back from settle- ments" in the Black Hills region. Inadc- (juate game protection laws were ineffec- tively enforced bv the state of South Dakota between 1871 and 1889 (Elley, 1945:5), and intensive trapping steadily reduced population hn els. According to Harris and Aldous (1946:348), the spe-
TURNER: MAMMALS OF THE BLACK HILLS
89
cics was nearly extinct in western South Dakota by 1927, but, reports of Seton (1929c: 453) and District Forester Allen S. Peck ( U. S. Biological Survey files) indicated tliat more than two tlionsand beaver inhabited the Black Hills National Forest at this time. In 1927, a nonpoliti- cal form of Game Commission was estab- lished ( Elley, loc. cit. ) , and responsible management practices were initiated by the South Dakota Department of Fishes, Game and Parks in the 1930's. Beaver were restocked in the Hills from popu- lated streams elsewhere in South Dakota. For example, due to the surplus of beaver along the James, Vermillion, and Sioux rivers in the southeastern part of the state in 1940, 300 of these large rodents were transplanted to the Hills (Wildlife Tips and Briefs, 1:14-15, 8 November 1940). Sale of 1196 beaver pelts netted trappers in the Black Hills $35,285.00 in 1945 ( Elley, he. cif. ) . Adequate protec- tion, in conjunction with the relatively low prices now being paid for beaver pelts, have allowed the species to in- crease in the Hills to the point that it now is overusing the habitat. In recent years, beaver have caused extensive dam- age to white birch, aspen, poplar, willow, and Cottonwood trees along many drain- age systems ( Sonth Dakota Conservation Digest, 24:12, October, 1957).
Dams of this aquatic mammal regu- late runoff by impounding water, thus aiding in the control of erosion and the maintenance of an adequate water table. Sediment gradually accumulates behind the dams and many former beaver ponds eventually are transformed into mead- ows, thereby increasing the stock carry- ing capacity of the range. Berner (1953b:9) noted that the three to four thousand beaver in the Black Hills Na- tional Forest, in combination with the grazing activities of cattle, have been, and still are, converting many brushy bottomlands into bluegrass pastureland.
Castor canadensis missouriensis Bailey Beaver
Castor canadensis missouriensis Bailey, 1919, Jour. Mamm., 1:32, 28 November (type
locality, Apple Creek, 7 mi E Bismarck, Burleigh Co., North Dakota).
The beaver is found on every major creek and river in South Dakota, l)ut it is most numerous in the Black Hills where liundreds of beaver dams decorate the countryside (Gilliland, 1968b:5). Morgan (1968:130) first reported this species from the study area when he ob- served that the numerous dams on streams of the Black Plills were unusu- ally small (in comparison to those he studied around Lake Superior), ranging from 50 to 100 feet in length and two to three feet in height.
Castor canadensis missouriensis dif- fers from C. c. concisor of south-central Wyoming in its slightly paler underparts, relatively broader zygomata, more open orbit, narrower nasals, and in the shape of the jugal (Long, 1965:622-623).
Some stocks of beaver have been in- troduced into the Black Hills from areas outside of South Dakota. For instance, three young from Yellowstone National Park were planted along Squaw [^Grace Coolidge] Creek in Custer State Park in 1914, and another familv of beaver was added in 1916 (South Dakota Conserva- tion Digest, 36:3, 1969). Sixty beaver were transplanted to the Hills from an unmentioned source in 1932; a pair on Jim Creek and another pair along Cold Springs Creek produced five and three young, respectively (reports of Regional Ranger, Black Hills National Forest, U. S. Biological Survey files). In 1947, five white beaver were observed along Grizzly Creek, a pair of which was cap- tured and placed in the Custer State Park Zoo (American Fur Breeders, 21:62, December, 1948; Game News of Penn- sylvania, 19:16, March, 1949).
From 1936 to 1944, 34 artificial sites, consisting of small dams and temporary bank lodges, were constnicted in the northern Black Hills ( Harris and Aldous, 1946:349). One hundred and thirty-six beaver were released at these artificial sites and 259 were released at natural sites. Members of the latter group tended to move on to other areas, whereas beaver became established on 31 of the
90
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
artificial sites. Of 100 beavers tagged and released in the period 1942 to 1944, 14 were retaken (op. c/f.:350); time be- tween release and recapture averaged 11.9 (1 to 33) months. One individual was obtained at the site of release; and the remaining 13 individuals moved an average of 9.1 (3 to 30) miles before being recaptured. The average rate of movement was 3.2 (0.3 to 14.0) miles per month. In all, 1892 beaver were trapped in 700 square miles of the north- em Black Hills from 1936 to 1945; in a five-day period in November 1944, 25 beaver were captured near the head of the South Fork of Boxelder Creek (loc. cit.).
Gilliland (1968b:5) noted that col- onies in the Black Hills consist of one or more families (one to 12 beavers) of all age groups (i. e., parents, kits, and year- lings). Two to eight kits usually are born between late April and early June.
Specimens examined (5). — SOUTH DA- KOTA: Lawrence County: Tinton, 2 mi N, 13 mi W Lead, 6000 ft, 1; 2 mi S Tinton, 6100 ft 1. Fennington County: Rapid Creek, 1..5 mi W Rochford, 5300 ft, 1 (UMMZ); Spring Creek, south of Rapid Cit>', 1 (MHM). Custer County: Palmer Gulch, 8 mi SE Hill City, 1 (FMNH).
Family CRICETIDAE— Nath^e
Rats, Mice, and Voles
This family includes a large assem- blage of terrestrial or partiallv fossorial rodents representing two subfamilies in the Black Hills. The more primitive cricetines are nocturnal in habit and are characterized by cheekteeth that bear two longitudinal rows of tubercles. The genus Neotoma is an exception; its teeth are cuspless, being formed instead of oc- clusal lakes of dentine surrounded by enamel. The more advanced microtines may be active for short periods through- out any given 24-hour period, usually following well established runways, and are characterized by rootless, prismatic cheekteeth that lack enamel on occlusal surfaces.
Three cricetine genera ( four species ) and three microtine genera ( five species )
in habit the Black Hills. The status of three additional cricetids (two cricetines and one microtine) in the Hills region remains uncertain, and another micro- tine has been incorrectly reported from the area.
As part of their reforestation program, the U. S. National Forest Service main- tains numerous ponderosa pine planta- tions throughout the Black Hills. These sites are prepared by windrowing, and the resulting dense grass and forb com- position that develops in these windrows provides excellent habitat for small mam- mals. Periodically, populations of micro- tines in these areas increase to levels that are not controlled by predators and tre- mendous damage to pine plantings re- sults. Such a population buildup of voles occurred in winter of 1968-1969, when an 870-acre area in the Bearlodge Dis- trict suffered 75 percent pine seedling mortality (Fred Wild, Range Conserva- tionist, pers. com.) and the Taylor Di- vide-Blacktail Creek plantation in the same district was nearly a total loss ( John C. Windsor, District Ranger, pers. com.).
Most damage occurred under the cover of snow; bark was removed from the root collar upward to the extent of the protective snow cover. Pine seedlings up to one inch in diameter frequently were gnawed completely through and other species such as aspen and oak were killed by girdling. Subsequent surveys of damaged areas yielded from two to 42 mice per 100 trap nights. For example, on the Dry Beaver Plantations, Penning- ton County, 20 Microtus penn.sylvanicus, two Clethrionomys p,apperi and two Sorex cinereus were obtained in 50 trap nights (Fred ^^^ild, pers. com.). The Forest Service plans to attempt control measures on all areas sampled that yielded 15 or more rodents per 100 trap nights (John C. Windsor, pers. com.).
Reithrodontomys megalotis dychei J. A. Allen
Western Harvest Mouse
Reithrodontomys dychei J. A. Allen, 1S95, Bull. Amer. Mus. Nat. Hist., 7:120, 21 May (type locality, Lawrence, Douglas Co., Kansas).
TURNER: MAMMALS OF THE BLACK HILLS
91
Reithwdontomijs megalotis dtjchei — A. H. How- ell, 1914, N. Amer. Fauna, 36:30, 5 June.
Reithwdontomijs megalotis dijchei is distributed throughout the Great Plains of central United States, inhabiting grassy environs, weedy borders of culti- vated tracts, thickets of shrubbery, and riparian communities. This mouse is ac- tive the year around, and in the Black Hills occurs at elevations up to 6200 feet, although it is most abundant in the tran- sition zone below 5400 feet. The first specimens of the western harvest mouse taken in the study area were obtained by B. Bailey at Spearfish early in August 1927, but R. meii,alotis was not reported from the Hills until much later, by Jones and Mursaloglu (1961:25).
This species is fairly common along the prairie border and through the foot- hills of the region. Nine individuals from southwest of Minnekahta were trapped in mid-June 1967 along fencerows of bluegrass, brome grass, cord grass, west- era wheat grass, and scattered forbs, and often were taken in runways of Microtus ochrogaster. The mixed-grass prairie lowlands that are interspersed with bushes and young trees in Wind Cave National Park also are inhabited, al- though to a lesser extent, by harvest mice in association with Etitamias minimus, Perognathiis fasciatus, P. hispidus, Pero- myscus manicidattis, M. ochrogaster, and Mils musciihis. In the Red Valley and foothills, moist draws and riparian com- munities that support growths of choke- cherry, wild plum, wild rose, hackberry, and other thickets of shmbbery provide adequate cover for harvest mice. For example, along the brushy streamside liabitat of Beaver Creek Canyon, 15 R. m. di/chei were live-trapped in late June 1968, along with 18 M. pennsylvanicus, 25 P. nuinicuJatus, 10 Zapus hudsonius, and one juvenile Neotoma cinerea. Five of these small rodents were collected near Hot Springs on 4 September 1968, on grassy ridges and in apple orchards that flank the Fall River.
At higher elevations, individuals were collected in erosional ditches, hayfields,
and along the interface of forested areas. J. A. King obtained two females in snap- traps placed under a log and beneath an oxerhanging creek bank in a spruce- filled canyon southeast of Hill City, and he took 12 additional harvest mice in a three- week period (28 December 1945 to 15 January 1946) from a gopher mound located in an upland pasture in the same general area.
Only eight of 56 R. megalotis from the Black Hills area that are housed in the Museum of Natural History are adult females, so reproductive data are meager. In 1968, harvest mice in Wind Cave Na- tional Park evidently bred from early spring to late summer. A pregnant fe- male was live-trapped and released in Beaver Creek Canyon on 27 June, and a lactating female was taken there on 2 July. Two pregnant females, one carry- ing two embryos that were 19 in crown- rump length and the other with four embryos (10 in crown-rump length), were obtained in the same canyon on 26 July. Two additional pregnant females, each carrying four embryos that were six in crown-rump length, were captured near the Park Headquarters on 30 Au- gust and 1 September. In other parts of the Hills, young of the year were ap- parent in late May to late August. Testes of 11 summer-taken males were 7.2 (6-8) in length.
Juvenal pelage of this mouse is short, fine and dark grayish-brown in color (Jones and Mursaloglu, 1961:14). Five adults from Fall River County taken in mid- June 1967 were undergoing the proc- ess of molt as evidenced by new hairs over the anterior three-quarters of the sides and dorsum, and by the line of de- marcation between fresh summer pelage and the worn winter pelage still present on the rumps of these individuals. Other specimens obtained at this time either were in fresh summer pelage or showed no signs of pelage replacement. Three individuals captured in Wind Cave Na- tional Park late in August 1968 were molting simultaneously o\'cr most parts of the body from the summer pelage to
92
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
the longer and more dense winter pelage.
Harvest mice from the Black Hills are parasitized externally by ticks, Ixodes spinipalpis Hadwen and Nuttall, fur- mites, Dennacarus hypudaei (Koch), laelaptid mites, AndrolaeJaps fahren- holzi (Berlese) and Omithomjssus ha- coti (Hirst), chiggers, Euschoengastia fasolla Brennan and Beck, fleas, Ma- laraeus telchinum (Rothschild) and MonopsyUiis wagneri (Baker), and an unidentified Anopluran louse that is rep- resented in collections only by the nymph stage.
The preponderance of males in col- lections from the Black Hills invalidates any attempts to test for secondary sexual dimorphism in R. megalotis. When study- ing harvest mice on the Great Plains, Jones and Mursaloglu (1961:12-13) found that the females are, on the aver- age, larger than males in all external and several cranial dimensions. However, in- dividual variation greatly exceeded sec- ondary sexual variation and statistically significant differences were lacking. A pattern of variation in coloration or size in the Hills was not readily discernible in the present study.
The remarkable uniformity in pelage color of R. megalotis over a considerable geographic range also was noted by Jones and Mursaloglu (1961:16). Most grassland cricetine species on the Great Plains have a dark eastern population and a pale western population. Environ- mental conditions along riparian habitats across the plains are somewhat similar to conditions of more eastern habitats, pos- sibly lessening differential selection pres- sures of R. megalotis in comparison to that incurred by nonriparian grassland cricetines (loc. cit.). No significant geo- graphic variation in color or size is evi- dent in specimens examined from areas surrounding the Hills in South Dakota and Wyoming (Table 15). Cranial meas- urements were taken in the manner de- scribed by Hooper (1952:9-11). Addi- tional external and cranial measurements (not included on Table 15) of 16 adults from the Hills are: tail length, 62.5±
9.47; hind foot length, 16.6±0.S1; ear length, 13.6±1.33; interorbital constric- tion, 3.2±;0.13; length of incisive fora- men, 4.4±0.15; depth of skull, 7.8±0.1S.
Specimens examined (96). — SOUTH DA- KOTA: Meade County: Black Hawk, 1 (AMNH). Pennington Countt/: Rapid City, 8 (SDMT); Moon, 22 mi W Hill Citv, 6200 ft, 1; 3-4 mi SE Hill City, 53-5400 ft, 22 (UMMZ); Spring Creek, 4 mi ENE Rocker- ville, 3600 ft, 1 (UMMZ). Custer County: Roby Canyon, 2 mi N, 22 mi W Custer, 5200 ft, 2; 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; 3 mi N Pringle, 1 (UMMZ); 5 mi N, 2 mi E Wind Cave, Wind Cave National Park, 4200 ft, 1; 1.5 mi N, 0.5 mi W Wind Cave, Wind Cave National Park, .3250 ft, 1; Beaver Creek Canyon, Wind Cave National Park, 4200 ft, 13; Headquarters, Wind Cave National Park, 4100 ft, 5; Wind Cave Canyon, Wind Ca\'e National Park, 4100 ft, 12; Wind Cax'e National Park, 5 (UMMZ); unspecified locality, 1 (UMMZ). Fall River County: 5.5 mi E Minnekahta, 4000 ft, 1; 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 9; 1 mi N, 5.5 mi E Hot Springs, 3400 ft, 2; 1 mi N, 6 mi E Hot Springs, 3400 ft, 3; Hot Springs, 1 (UMMZ); 4 mi E Hot Springs, 3400 ft, 1; 2 mi S, 2 mi E Hot Springs, 3600 ft, 1.
WYOMING: Crook County: 1.5 mi NW Sundance, 5000 ft, 3.
Additional records.— SOUTH DAKOTA: Lawrence County: Spearfish (BB).
Peromyscus leucopus aridulus Osgood
White-footed Mouse
Peromyscus leucopus aridulus Osgood, 1909, N. Amer. Fauna, 28:122, 17 April (type locality, Fort Custer, Big Horn Co., Mon- tana).
The white-footed mouse occurs throughout the northeastern two-thirds of North America. The distribution of Feromyscus leucopus across the arid Great Plains closely approximates the dendritic pattern of mesic deciduous communities that follow major drainage systems. The species is relatively un- common in the Black Hills. It inhabits the notably discontinuous deciduous woodland formations up to 6100 feet, although it is more abundant below 5000 feet. No specimens are at hand from Fall Bivcr County, but white-footed mice probably occur there. Although this mouse has been taken in the Hills only
TURNER: MAMMALS OK 11 IK BLACK HILLS
93
o
2
o
a P
B
en CS HI
c
a;
o ■«
o
CD
e o
O
>
a
o
OJ
lO
|
AVOJ-mOOJ |
o CO '—1 |
^ o |
coS |
00 C<1 o |
o CO — 1 |
|
A.1B[[I\-BUI |
CO d |
CO d |
CO CD |
CO CD |
CO d |
|
JO i[]Suaq |
+1 |
+1 |
+1 |
+1 |
+1 |
ippwajq
3I4BUloSX2
l[n>is JO
iqSFA\
CO
o
+1
aniq
U33.lS
p9J
;uaoa9j
ceo
.^ cj c8 ^ ^ QJ
3 C
CM
CO
Q
CO
|
Of |
-^ ^ |
^^ |
Of |
^^ |
|
CM |
o+ |
Of |
Oi ^ |
Of |
|
^ c |
o c |
^ c |
. G |
TP C |
|
^ gQ |
-2Q |
^ 2Q |
03 |
|
|
3S- |
CO '^ |
CO "^ |
^S" |
|
|
^— ' |
— ■ |
|
■ — ■ |
^— ' |
|
CD |
CO |
-^ |
CD |
00 |
|
1— ( |
CM |
Q
C/2
C30
CO d +1
|
CO CO |
<M |
—1 CO |
05 (M CO |
o CO ^ |
||
|
j> d +1 |
t-^ d +1 |
i> d +1 |
t-^ d +1 |
t-' d +1 |
t-^ d +1 |
i> d +1 |
|
i-H |
^H |
05 |
CM |
■'-M |
CM |
05 |
||||||||||
|
t- "^ |
c:5 |
CO |
in |
CO |
00 |
I-H |
q |
C>1 |
00 |
CO |
O (M |
|||||
|
d d |
d |
d |
d |
d |
d d |
^H |
d |
d |
d |
I-H d |
||||||
|
1— 1 |
+ 1 |
1— 1 |
+1 |
^H 4-1 |
II |
iS |
i-H |
+1 |
I-H |
+1 |
I-H |
II |
^ II |
|||
|
^ |
t^ |
o |
CD |
o |
I-H |
« |
t^ |
CD |
bO C 00 |
|||||||
|
I— 1 |
IC |
CO |
CO |
qS |
CD |
I-H |
CD |
o |
05 |
TP |
CO |
t- |
•- TP in |
|||
|
tc^ |
d |
I-H |
d |
d |
I-H |
d |
rM |
d |
d |
I — 1 |
d |
c — ; d |
||||
|
+1 |
(M |
+1 |
^ CM 3 |
+1 |
Ol |
+1 |
Q |
(M |
+1 |
CM |
II |
5.^^ II |
||||
|
o |
CS |
O |
Q |
r- |
> |
|||||||||||
|
>. ^ |
4^ o |
CD 6 |
C/D d |
c/5 |
bO |
|||||||||||
|
15 |
^ |
Q |
CM |
U |
00 |
u |
(M |
I-H |
G |
i> |
C CO |
|||||
|
C C5 |
i-H |
1—" |
O oq |
CO I-H |
be CM |
1 — 1 1-H |
C |
q |
(M CM |
u |
CM CO |
I-H CM |
O c o |
in CM |
CM co" |
O CM ^ |
|
Dakota 1 |
+ 1 |
3 O C/3 >> C |
I-H |
+ 1 |
ashabau 1 |
+1 |
> fa |
I-H |
+ 1 |
4-1 4-1 cu PQ |
I-H |
II |
I-H |
II |
2^ +1 O |
|
|
J5 -1— > |
in |
o |
CO |
^ |
o |
-t-T C O |
CO |
O |
U |
o |
^ CM |
|||||
|
c)5 ^ |
CO CO |
1> |
in d |
alley, 143.0 |
o c^i |
CD |
o 00 |
o in |
in -p |
"^ |
d |
CD |
e, and 140.6 ±9.6 |
|||
|
CO ^"r— 1 |
+1 |
CO I-H |
+1 |
+1 |
be |
CO I-H |
+1 |
CO I-H |
II |
"E |
-p I-H |
II |
||||
|
> bc |
13 |
o |
ca u |
|||||||||||||
|
o |
1— 1 |
CM |
c o 1-J |
t- |
c C8 |
'^ |
4-1 |
CO |
O |
|||||||
|
"^ CO |
CM d |
05 in |
CO d |
o d |
CO O |
in |
I-H d |
as |
(M d |
CO |
I-H d |
i; CM o |
||||
|
CM |
+1 |
(M |
+1 |
■g<M |
+1 |
CM |
+1 |
O C/2 |
(M |
II |
(M |
II |
Ji"^' II |
|||
|
O CM |
O |
|||||||||||||||
|
CM |
CD |
o |
CM |
<M |
CO |
l^ |
||||||||||
|
I> |
I-H |
CO |
r-{ |
CO |
(M |
05 |
I-H |
CM |
I-H |
'^ |
I-H |
I-H ^^ |
||||
|
t-' |
d |
CD |
d |
t-^ |
d |
CD |
d |
i> d |
t-' |
d |
t~^ d |
|||||
|
CM |
+1 |
CM |
+1 |
CM |
+1 |
(M |
+1 |
CM |
II |
CM |
II |
CM II |
|
CM CO CM |
CO ^ CM |
CO in ^ |
in I-H I-H |
CO 00 CM |
I-H I-H |
a> O I-H |
|
00 d -^ II |
i> d ^ II |
00 d ^ II |
00 d ^ II |
t-^ d -* II |
d d ^ II |
d d ini, |
|
.I0[00 JO 9U0X |
CM O |
^ 1> |
CO CM Gi |
CD |
00 CO q |
CO t-; in |
co2 |
|
l-^ CO <M II |
t-^ CO CM II |
CD oi (M II |
d cm' CM II |
d oi <M II |
CM II |
d CO CM II |
Of
00
Q
C/3
94
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
from mid-June to mid- August, it is active the year around and does not hibernate. J. A. Allen (1894a: 325) first reported white-footed mice from the Hills under the name Sitomys americamis arcticus (see below).
Big Spearfish Canyon extends from Savoy, where the streambed elevation is about 5000 feet, northward, emerging from the Black Hills at Spearfish (3600 feet). Spearfish Creek courses below massive clifi^s, with luxuriant vegetation ( bur oak, American elm, boxelder, green ash, and deciduous shrubbery) on the canyon floor. Sixteen ( 11 males and five females) P. leucopus were taken in the underbrush of this riparian woodland in early August 1967. Vanocker Canyon arises on the southeast slopes of Dead- man Mountain (4800 feet) and extends northeastward, emerging at Sturgis ( 3600 feet). Five individuals (four males and one female) were obtained in this can- yon on 28 June 1967, and two more ( one of each sex) were collected there on 15 August 1967. These mice were trapped in brush in a pasture that bordered an aspen grove, and in a mixed oak, pine, and aspen woodland.
Nine Pewmysciis leucopus ( five males and four females) were captured in the Central Basin, northeast of Custer, in late June 1967. These were taken in aspen woodlands bordering Iron and Grizzly creeks; the woodland contained much undergrowth, fallen logs, decid- uous shrubbery and rock outcroppings. Eutamias minimus, Peromyscus manicu- latus, Clethrionomys gapperi, Microtus longicaudus, and M. pennsylvanicus are associates of P. leucopus in the Hills.
White-footed mice also inhabit allu- vial wooded bottomlands of drainages in the Red Valley and foothills. Specimens collected along Squaw Creek in late July 1894 by W. W. Granger were taken in an area that, "is wooded with aspen, willows, boxelders and other deciduous trees" (J. A. Allen, 1895a: 261). Granger also captured eight P. leucopus well out on the prairie east of the Black Hills, about seven miles southwest of the mouth
of Spring Creek. This site is mentioned as a further example of the restrictt^d habitat of P. leucopus on the plains. Spring Creek was "bordered by box- elders, cottonwoods, plum thickets, wil- lows, wild currants, and rank weeds and grass" at the time of Granger's visit ( loc. cit. ) .
Throughout its range in the Black Hills, the white-footed mouse is sym- patric with the deer mouse. These two species are often taken side-by-side in the same trapline (see account of Pero- myscus maniculatus) and are frequently difficult to difi^erentiate. For example, J. A. Allen (1895a: 268) listed 39 speci- mens of Peromyscus leucopus from Cus- ter and 20 from Squaw Creek. He noted that a few specimens presented a decided fulvous or reddish wash, and that one specimen ( AMNH 9370) approached the characteristic "fulvous tint of nehracen- sis" [P. maniculatus], Allen later (1899: 15) assigned all of these specimens to a new subspecies, '^Peromyscus texanus suharcticus [P. m. artemisiae]," stating "Black Hills specimens are not at all typical, some of them decidedly ap- proaching P. t. nehracensis, while others are quite like the Montana and Saskatch- ewan examples." Only 10 of the 59 speci- mens listed by Allen are actually P. leu- copus, the remainder being maniculatus (including AMNH 9370). Externally, white-footed mice are larger in body size, have longer tails and hind feet, and the summer pelage is somewhat duller, with a less distinctly bicolored tail, in comparison with deer mice. Subadults of the former species are easily confused externally with adults of the latter, but the larger, more robust skull of P. leu- copus, coupled with age determination by degree of tooth wear, readily resolves this dilemma.
Three of six adult female P. leucopus from the Hills, for which reproductive data are available, showed signs of repro- ductive activity. One taken northeast of Custer on 26 June 1967 carried six em- bryos that were 3 in crown-rump length.
TURNER: MAMMALS OF THE BLACK HILLS
95
Another, obtained southwest of Sturgis on 28 June 1967, was lactating and had five placental scars present in the repro- ductive tract. The third, captured south of Sturgis on 15 August 1967, had four placental scars. The average length of testes of 11 adult males collected in the summer was 11.8 (10-13).
Many adult white-footed mice taken in late June and early July were molting over the dorsum and sides, whereas other individuals already were in fresh summer pelage. Juveniles have soft gray fur. Two subadults from northeast of Custer ( 19 June 1967 ) were undergoing post-juvenal molt in which new brown hairs were emerging along the sides, whereas the dorsum still was furred in the grayish juvenal pelage.
Examination of 26 P. leucopiis (16 males and 10 females) from the Black
Hills revealed no apparent secondary sexual variation in coloration or in ex- ternal and cranial measurements, nor was a pattern of variation in these param- eters discernible in tlie Hills populations.
Specimens from the Black Hills, and adjacent areas to the west in Crook County, Wyoming, are darker in average tone of color, and much smaller in most external and cranial dimensions when compared to populations on the South Dakotan plains (Tables 16 and 17). The small size of the Black Hills representa- tives has been a source of past mis- identifications.
Feromijscus leucopiis probably ex- tended its range westward on the Great Plains along major drainage systems in mesic Post-Wisconsin times. The pres- ently isolated segments formerly were part of a more or less continuous and
Table 16. — Geographical variation in coloration of the mid-dorsal pelage of adult Pewmyscus leucopiis obtained in summer from the
Northern Great Plains.
|
Number and sex of specimens averaged |
Tone of color |
Percent red reflectance Percent green reflectance |
Percent blue reflectance |
|
26 (16,$, 10 9 ) Mean SD |
Black Hills, South Dakota and Wyoming 26.7 48.4 27.4 24.2 ±3.60 ±0.28 ±0.17 ±0.17 |
||
|
6(5cJ,19) Mean SD |
26.2 ±1.37 |
Crook County, Wyoming 48.4 28.3 ±0.22 ±0.20 |
23.3 ±0.12 |
|
5(45,19) Mean SD |
Washabaugh County, South Dako 26.4 48.5 27.0 ±2.86 ±0.22 ±0.17 |
ta 24.5 ±0.26 |
|
|
15(75,89) Mean SD |
26.9 ±2.47 |
Todd County, South Dakota 47.3 27.5 ±0.31 ±0.20 |
25.2 ±0.24 |
|
7(35,49) Mean SD |
27.4 ±2.39 |
Harding County, South Dakota 49.5 27.0 ±0.32 ±0.16 |
23.5 ±0.21 |
|
17 (75,109) Mean SD |
27.5 ±3.25 |
Bennett County, South Dakota 49.5 26.4 ±0.37 ±0.22 |
24.1 ±0.19 |
|
22 (115, 119 ) Mean SD |
30.1 ±3.26 |
Cherry County, Nebraska 49.5 26.6 ±0.26 ±0.18 |
23.9 ±0.17 |
96
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Table 17. — Geographic variation in selected external and cranial measurements of adult Peromijscus
leucopus from the Northern Great Plains.
|
Number and sex j- of specimens B m averaged [5 Ji |
Tail length |
Hind foot length |
Ear length Greatest length of skull Zygomatic breadth |
Rostral length |
Length of incisive foramen |
Skull depth |
|
|
38(25 5,13?) Mean SD |
164.2 ±12.98 |
70.6 ±10.72 |
Black Hills, South Dakota and Wyoming 20.7 16.5 26.4 13.6 9.8 ±0.33 ±1.29 ±0.82 ±0..54 ±0.49 |
5.4 ±0.25 |
9.6 ±0.28 |
||
|
6(55,1?) Mean SD |
171.5 ±13.69 |
75.3 ±5.95 |
21.8 ±0.75 |
Crook County, Wyoming 16.5 26.2 13.7 ±1.52 ±1.19 ±0.67 |
9.5 ±0.75 |
5.3 ±0.43 |
9.5 ±0.21 |
|
5(45,1?) Mean SD |
175.4 ±11.10 |
77.2 ±4.32 |
Washalsaugh County, South Dakota 21.4 17.0 27.2 14.1 10.2 ±0.55 ±1.22 ±0.84 ±0.56 ±0.58 |
5.5 ±0.27 |
9.8 ±0.18 |
||
|
16(85,8?) Mean SD |
172.2 ±13.41 |
74.9 ±6.51 |
21.8 ±0.75 |
Todd County, South Dakota 17.1 27.2 14.1 ±1.23 ±1.10 ±0.63 |
10.2 ±0.69 |
5.6 ±0.32 |
9.8 ±0.25 |
|
7(35,4?) Mean SD |
184.4 ±8.42 |
77.4 ±6.50 |
21.1 ±0.61 |
Harding County, South Dakota 16.7 27.7 14.7 ±0.76 ±0.45 ±0.32 |
10.5 ±0.28 |
5.5 ±0.23 |
10.0 ±0.22 |
|
17(75,10?) Mean SD |
176.3 ±9.95 |
77.8 ±8.59 |
21.3 ±0.86 |
Bennett County, South Dakota 16.0 27.2 14.4 ±1.62 ±0.97 ±0.61 |
10.3 ±0.54 |
5.5 ±0.28 |
9.9 ±0.32 |
|
34 (20 5,14? ) Mean SD |
170.1 ±7.76 |
72.9 ±4.57 |
22.0 ±0.65 |
Cherry County, Nebraska 15.8 27.2 14.4 ±0.73 ±0.73 ±0.51 |
10.3 ±0.48 |
5.5 ±0.23 |
9.8 ±0.25 |
interbreeding population (Jones, 1964: 197). Currently, gene-flow between these limited populations probably is re- duced or absent, and tliese populations presumably now are adapting to en- vironmental conditions independently.
Measurements additional to those on table 17 of 38 white-footed mice from the Hills are: weight, 23.4 ±4.66; inter- orbital constriction, 4.2 ±0.14; and length of maxillary tooth-row, 4.0 ±0.15.
Specimens examined (62). — SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- yon, 1.5 mi S, 0.5 mi E Spearfish, 3800 ft, 7; Big Spearfish Canyon, 3 mi S Speai-fish, 4200 ft, 6; Big Spearfish Canyon, 5 mi S, 2 mi W Spearfish, 4600 ft, 1; Big Spearfish Canyon, 9 mi S, 3 mi W Spearfish, 5000 ft, 2; 2 mi S Tinton, 6100 ft, 1; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 1; 2 mi W Nemo, 4700 ft, 2. Meade County: Vanocker Canyon, 2 mi S Sturgis, 2; Vanocker Canyon, 3 mi S, 0.5 mi W Stiugis, 4400 ft, 5. Pennington County: 1 mi S, 8.5 mi W Rapid City, 1. Custer County: 5.75 mi N, 5.75 mi E Custer,
5220 ft, 9; Custer, 3 (1 USNM, 2 AMNH); Custer State Park, 1 (UMMZ); Squaw [Grace Coohdge] Creek, 3000 ft, 8 (7 AMNH, 1 FMNH); 0.25 mi N Otis, 2 (UMMZ).
WYOMING: Crook County: 15 mi ENE Sundance, 3825 ft, 9; Sundance, 2 (USNM).
Additional record.—SOVTK DAKOTA: Lawrence County: Spearfish (BB).
Peromyscus maniculatus nebrascensis
(Coues)
Deer Mouse
Hesperomys sonoriensis var. nchrasccnsis Coues, 1877, in Coues and Allen, Bull. U. S. Geol. Surv. Territories, 11:79, August (type lo- cality restricted to Deer Creek, approxi- mately fi\e miles from its mouth, Converse Co., Wyoming, by Jones, Proc. Biol. Soc. Washington, 71:108, 16 July 1958).
Perom i/sctis maniculatus nebrascensis — Osgood, 1909, N. Amer. Fauna, 28:75, 17 April
In the Black Hills, this ubiquitous cricetid is by far the most abundant and widely distributed of the mammalian
TURNER: MAMMALS OF THE BLACK HILLS
97
species. Although it is most common in grassy cn\iions of the Red Valley and foothills, P. inaniciilatus occurs in a wide variety of habitats, including cultivated cropland, grazed or ungrazed pasture- land, grassland meadows, marsh and stream borders, spruce and aspen wood- lands, dense pine forests, and even among bare rock at the summit of Harney Peak (7240 feet), atop Devils Tower, and several hundred feet below ground surface in Wind Cave.
Dice (1939:5) estimated that the to- tal population of P. maniculatus in the Black Hills in late summer of 1935 was between one and five million individuals. He indicated that deer mice occurred in the greatest abundance in dense and medium dense stands of western yellow [ponderosa] pine in densities of 932 and 894 mice per square mile, respectively {op. cit.:2. Table 1). However, Dice ( 1939 : 1 ) based his estimates on only 39 specimens and erroneously assumed that the Black Hills population was isolated to a large extent. Due to poor trapping results in grasslands near Rapid City and south of Hot Springs, Dice did not trap in grassland, pasture, or cultivated fields in other parts of the Hills (op. cit.:3) as- suming that no deer mice occurred in these areas (op. cit.A).
My own investigations indicate that, although deer mice are omnipresent in the Black Hills, the highest population densities are attained in those environs dominated by graminoid vegetation. For example, in a five-day period ( 13-18 June 1967) 82 deer mice were captured near Minnekahta in arid pastureland of the Red Valley ( see account of Microtus ochwgaster for a description of the area) . In approximately 2000 trap-nights in the summer of 1968, 315 P. nuiniculatus were taken in the grassy uplands and semi-grassy canyon systems of Wind Cave National Park. On 27 August alone, 66 deer mice were captured in 150 trap- nights. Most of these specimens were discarded after being examined for ecto- parasites and autopsied for reproductive data. Because quadrat sampling meth-
ods were not used in the current study, calculations of population size were not attempted, but, it is certain that Dice's computations underestimated the num- ber of deer mice in the Black Hills.
In the northern foothills and canyons, and along some drainages in the central section of the Black Hills, P. m. nehras- censis is ecologically sympatric with P. leucopus (see account of that species). I have taken both species within a few feet of each other, and even in the same trap, in Little Spearfish Canyon. How- ever, P. leucopus is less common and is stenoecious, generally associating with elements of the eastern deciduous forest. Due to its occurrence in all available habitats, the deer mouse is closely associ- ated with each of the other terrestrial mammals in the Hills region.
Deer mice captured in Wind Cave National Park in summer of 1968 were grouped into age classes based on tooth wear, general dimensions, and type of pelage. Of 315 individuals, there were 188 adults, 116 subadults, and 11 juve- niles. Mice taken in July (97) and Au- gust (203) were further segregated by sex. In the following analysis of popu- lation composition, percentages for July are followed by those for August: adult males, 32.0, 32.0; adult females, 25.8, 28.1; subadult males, 19.6, 12.8; subadult females, 20.6, 22.7; juvenile males, 1.0, 3.4; juvenile females, 1.0, 1.0; The analy- sis indicated that about 40 percent of summer populations of P. maniculatus in the Black Hills are immature individuals.
This species may breed most months of the year in the Hills region, but ade- quate data are available only for the summer. J. A. King captured two preg- nant females southeast of Hill City, one of which carried five embryos ( 14 March 1946) measuring three in crown-rump length, and the other containing two embryos (25 March 1946) measuring 23 in crown-rump length. On 5 April 1946, King obtained two gra\'id females north of Newcastle that each carried five em- bryos which had an average crown- rump length of 2 and 3, respectively.
98
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Of 257 female P. nianiculatus ob- tained in the summer from the Black Hills and autopsied, 48.3 percent showed indications of reproductive activity. Be- cause many nearly-mature subadults car- ried fetuses, both adults and subadults were examined in this study. Of 48 fe- males collected in June, 16 were preg- nant, having an average of 5.2 (4-7) embryos that were 9.3 (4-19) in crown- rump length. Four placental scars were manifest in the reproductive tract of another individual, and seven females were lactating. The average length of testes of 67 males obtained in June was 9.0 ( 6-18) . In July, 24 of 87 females were parturient, and carried 5.0 (4-9) fetuses; the average crown-rump length of the latter was 11.8 (2-25). Three other indi- viduals had 4.3 (4-5) placental scars; a like number of females had swollen uteri, possibly indicating recent implantation. Twenty additional females were lactat- ing. The average testicular length of 64 males taken in July was 9.6 (6-18). Of 97 females obtained between 16 and 31 August, only 11 were gravid, containing 4.6 (4-6) embryos with an average crown-rump length of 16.8 (6-27). Twelve other females had 4.3 (3-6) pla- cental scars, and nine other individuals were lactating. Six females had swollen uteri. Seventy-two males captured in the same period had an average testic- ular length of 9.8 (7-17). In the first week of September, six of 25 females contained 4.3 (3-6) young each that had an average crown-rump length of 16.8 (8-23). Three females had 5.7 (5-6) placental scars, and three other individuals were lactating. The aver- age testicular length of 27 males ob- tained in the period 1 to 7 September was 9.4 (5-18). Analysis of reproductive data on a monthly basis masks short- termed fluctuations. For example, the percentages of pregnant individuals en- countered over four successive two-week periods were: 1 to 15 June, 40.0 percent of 25 females; 16 to 30 June, 26.1 percent of 23 females; 1 to 15 July, 36.6 percent
of 41 females; and 16 to 31 July, 19.6 percent of 46 females.
Adults molting from winter to sum- mer pelage were taken as early as 10 June, but maximal time of pelage re- placement occurred from late June to mid-July. Osgood (1909:19) indicated that there is but one complete annual change in pelage of adult Peromyscus and that the process commences near the head, progressing caudad. Molting indi- viduals trapped in late August and early September appeared to be acquiring the characteristic long, dense pelage of win- ter, and apparently were undergoing a second seasonal molt. The high repro- ductive rate of deer mice results in many individuals of different ages occurring simultaneously in the same area through- out warmer months, thus complicating interpretation of seasonal aspects of molt. Subadults undergoing post-juvenal molt were observed in all periods. The slate- gray Juvenal pelage is replaced by brownish to buffy haii's that first occur along the upper portion on each side.
Twenty-two kinds of ectoparasites were harbored by P. maniculatus from the Black Hills: two lice, Hoplopleura hesperomijdis (Osborn) and Polyplax auricularis Kellogg and Ferris; three chiggers, Euschoengastia setosa (Ew- ing), Psuedoschoengastia farneri Lipov- sky, and Xenacarus plnmosus (Green- berg); four ticks, Dermacentor aiulersoni Stiles, Ixodes kingi Bishopp, /. spinipalpis Hadwen and Nuttall, and individuals of the /. ochotonae-angustus complex; six mites, Androhelaps fahrenholzi (Ber- lese), Dermacarus hypudaei (Koch), En- laelaps stahidaris (Koch), Laelaps mi- croti (Ewing), Oniitlionysstis bacoti (Hirst), and members of the genus Hirs- tiomjssus; and seven fleas, Monopsyllus wagneri (Baker), M. eumolpi (Roths- child), Mahraeus telchinum (Roths- child), Orchopeas leucopus (Baker), O. sexdentatus (Baker), CotaUagia decip- iens (Rothschild), and Epitedia icen- manni ( Rothschild ) .
Comparison of 56 male and 42 female deer mice from the Black Hills revealed
TURNER: MAMMALS OF THE BLACK HILLS
99
no significant secondary sexual dimor- phism in external or cranial measure- ments, although females are slightly larger in most dimensions. Males are paler in tone (29.8±4.12) and greater in reflectance of reds (50.6±0.36) than are females (27.4±4.85 and 49.5 + 0.28, re- spectively); however, there is consider- able variability in each of these param- eters.
Peromijscus maniculaius is quite vari- able in color and in external and cranial dimensions in the Black Hills; this is probably due to a greater divergence in topography, habitat, and edaphic factors in the Hills than in the surrounding prairie. Dice (1941:16) noted a general tendency for this species to increase in size westwardly over the Great Plains, and indicated (1942:9) that the presence of the Black Hills does not affect this trend in any significant manner. Deer mice in the Hills are intermediate in size between eastern, northern, and western populations, but are assignable to the western subspecies (Table 18). Pero- mijscus manicidatus nebrascensis differs from P. m. hiteus to the east in being larger externally and cranially, and in being darker dorsally ( Fig. 13 and Table 19). Osgood (1908:78) noted that, al- though typical nebrascensis average de- cidedly larger than typical liiteus, indi- vidual variation in each race is great enough to allow overlap in the range of various parameters. For example, he re- garded five of 13 specimens from Elk Mountain as resembling Ititeus, and the other eight as resembling nebrascensis. Occasional high frequencies of paler and smaller mice in the foothills and periph- eral areas may be expected due to gene exchange with populations of the sur- rounding plains. Specimens from Rapid City, for example, approach the dimen- sions of luteus rather than nebrascensis.
Stebler (1939:391-392) and Dice (1942:1-10) both discussed the correla- tion between soil color and pelage color of P. maniculaius in the study area. In general, the soils of the Black Hills are dark in color, varying from reddish to
blackish brown. When comparing habi- tats of the l^adlands with those of the Black Hills, Dice (1942:6) stated: "On the forested Black Hills, on the contrary, the variation from place to place in soil color is relati\ cly slight, and there is only a small amount of variability in the color of deer-mice." In actuality, soils of the Hills are exceedingly diverse in colora- tion. For example, the two soils col- lected south of Spearfish (Table 20) were within 20 yards of one another; sample A was obtained mid-way up a steep slope, and sample B was taken at the base of the slope. Soils from near Rapid City register abnormally pale in tone due to a high gypsum content that increases reflection properties of the soil. This great diversity in coloration of soils invalidates comprehensive correlations between pelage and soil hues, unless a soil sample is taken at each site of cap- ture throughout the entire Hills region. Collectively, soils of the Red Valley "racetrack" differ significantly from those of the Limestone Plateau and Cen- tral Basin, being paler in tone and re- flecting reds more intensely. Soils of the Central Basin average somewhat paler in tone than those of the Limestone Pla- teau, but otherwise they are fairly simi- lar.
Peromijscus manicuJatus in the Black Hills is much darker on the average than either the small "ochraceous-buff" race on the arid plains south and east of the Hills, or the large "reddish" race on the sandy soils of eastern Wyoming ( Fig. 13 and Table 19). Collectively, specimens from the southern Hills (especially from southwest of Minnekahta) are, on the average, paler than those from the north- ern section, and specimens from the western part (especially from the vicin- it>^ of Ditch Creek, Beaver Creek, Four Corners, and Buckhorn) have greater reflection of reds than individuals from the eastern Hills.
Measurements additional to those given in Table 18 for 98 adult specimens from the Black Hills are: zygomatic
100
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
CO
C
t8
o
■s
o J3
o
CO
5
3 C
CO
8
3 OS
(8
C
C G
s o
•iH
ca V a >
:a
a
o O
00
PQ
AiE[[I.\-BUI
uauiBJOj
aATspuT
JO q:(Su9q
ipdap I[iMS
[Biisoy
ipgU9[
4sa;BajQ
ni§PM
i{:|§ua[
1{4§U8|
looj pujH
q}gua[
q}Sua[
|
CD —1 |
in 1> I-; |
CO |
|
|
CO o +1 |
00 o +1 |
CO O +1 |
CO o +1 |
in
CD
C5 cq
00 o
+1
|
CO |
00 CO cq |
cq I— 1 |
CO cq |
|
o +1 |
in d +1 |
ic o +1 |
+1 |
o
a Q
G o
o U
c
c
CO
-^
o
+1
Oi CD
d +1
« cq
Co
O CO ' CO
+1
CD lO O
05
ai d +1
> 1> CO "^05 0 -^ +1
in I— I oq
ai d
+1
C8
^t-: CO
^05 0
o a Q
00
C2 CO
'^ d
O
^ 00 O iS CO -LI *\
s <»
^ lO
+1
CO
d +1
G O
3 m c oq U
C
45 CO
Ph CD lO
co-^oq CO l'^^ +1
S
to t>
S .-H -^
~ CD ^
c c:5 d
S +1
o
^ 00 rr
S d d
o
n3 .
+1
d +1
liO i>
■CO CO
" +1
cq
> a o
U
|
-^ +1 |
a: |
1 — i |
+1 |
ft.; |
^ +1 |
5j C |
|
f— ( |
1—1 |
tr- |
S |
|||
|
CO CD |
1—1 |
q |
<j> 00 |
|||
|
00 r-^ |
d |
1 — ( |
00 d |
ftn' |
||
|
^ +1 |
1—1 |
+1 |
-" +1 |
CO
00 in t-^ 1-! ^ +1
in o
Ci
+1
|
cq o |
o t* cq |
in in |
CO ^ o |
|
d in CO II |
rA CD CO II |
1-! in CO II |
CD II |
|
05 CO in |
CD cq in |
p cq |
cog |
|
t-^ 00 3+1 |
d CD in 1—1 ^ +1 |
cq d in 1—1 ^ +1 |
in .-1 ^+1 |
|
T3 0) i-g |
^-^ |
y-~. |
.—v |
^-~, |
|
Of |
Of |
Of |
Of |
|
|
1—1 |
cq |
o |
cq |
|
|
r1 f-i Q^ rt G he |
1—1 |
^ |
^„c |
*^. c |
|
Iinber ; f speci avera |
^ SQ |
^" Iq |
^ go |
^ s |
|
cq 5 <^ oq >< |
CD •^ C/^ in "^ |
s:^^ |
CO '^ |
|
|
— - |
- — ' |
— ' |
— - |
|
|
P o |
CO |
00 |
CO |
CD |
|
CO |
as |
Cl |
in |
Q
TURNER: MAMMALS OF THE BLACK HILLS
101
|
P M NEBRASCENaS |
> P M NEBRASCENSIS |
P M NEBRASCENSIS |
P M LUTEUS |
|||
|
BLACK HILLS |
HARDING CO, |
S |
D |
EASTERN WYOMING |
BENNETT CO . |
|
|
TONE |
282*4 58 |
304*2.93 |
331*5 15 |
348*457 |
||
|
%Reo |
49 8*0 54 |
30 7«0 2S |
910*0 25 |
48 2*0 30 |
||
|
%«IEEN |
26 9*019 |
25.3*021 |
26 1 *0 16 |
278*0 19 |
||
|
%SLUE |
23 7*019 |
24 0*011 |
22 9*014 |
240*0 19 |
Fig. 13. Geographic variation in coloration of the mid-dorsal pelage of four groups of adult deer mice collected during the summer. Pewmijscus maniculatus nehrascensis a\erages much darker in tone than the "ochraceous-buff" race, P. m. hiteus, to the south and east, and specimens of P. m. nehrascensis from the Black Hills average much darker than either the dark form from Harding County, South Dakota, or the "reddish" form that inhabits sandy soils of eastern Wyoming.
breadth, 13.1: striction, 3.9 d
t0.40; and interorbital con- 0.26.
Specimens examined (778). — SOUTH DA- KOTA: Lawrence Coiinhj: Big Spearfish Can- yon, 5 mi S, 2 mi W Spearfish, 4600 ft, 2; Big Spearfish Canyon, 6 mi S, 2 mi W Spearfish, 4600 ft, 1; Tinton, 5900 ft, 21; 2 mi S Tinton, 6100 ft, 37; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 1; Little Spearfish Canvon, Savoy, 4 (USNM); Deadwood, 1 (USNM); Roubiax Lake, 2 (FMNH); Du- mont, 1 (USNM); 2 mi W Nemo, 4700 ft, 6; 1 mi E Nemo, 4700 ft, 2; 3 mi E Nemo, 4650 ft, 1; Boxelder Creek Canyon, 2 (NRW); Steamboat Rock Campground, 3 (NRW); Jim Creek, east Merritt, 1 ( NRW ) . Meade County: 3 mi S, 0.5 mi W Sturgis, 4400 ft, 1; 6 mi S, 1.5 mi W Sturgis, 4600 ft, 2; Black Hawk, 9 (AMNH). Pennington County: 1.5 mi W Rochford, 1 (UMMZ); south slope Norris Peak, 1 (NRW); Castle Creek, 6500 ft, 1 (UMMZ);
Beaver Creek, 4 mi 6400 ft, 56; Beaver Deerfield, 6500 ft, 8
Bald Hills, Pactola, 1 (USNM); 8 mi NE Rapid City, 3 (UMMZ); 4 mi N Rapid City, 7 (AMNH); 3 mi N Rapid City, 3 (NRW); 2 mi N, 3 mi E Rapid City, 2; Rapid City, 65 (3 UMMZ, 3 USNM, 48 NRW, 3 SDMT); Dark Canyon, 1 mi S, 4 mi W Rapid City, 15 (AMNH); 4 mi SE Rapid City, 2 (UMMZ); N, 10.5 mi W Deerfield, Creek, 3 mi N, 9 mi W 3 mi N, 7 mi W Deer- field, 6900 ft, 3; Redfern, 9 (USNM); Rocky Mountain Range E.xperimental Station, McVey Bum, 2 (NRW); NE Sheridan Lake, 2 (NRW); Spring Creek Canyon, 3 (NRW); Wild Irish- man Gulch, 2 (NRW); 3 mi S, 1 mi W Rock- erville, 6; Spring Creek, south Rapid City, 6 (MHM); 4 mi NW Hill City, 4 (UMMZ); Moon, 22 mi W Hill City, 6200 ft, 4; Ditch Creek, 14 mi W Hill Cit>', 6400 ft, 22; 3-4 mi SE Hill City, 5300-5400 ft, 38 (UMMZ); 5 mi SE Hill City, 1 (UMMZ); 2 mi W Oreville, 5500 ft, 1 (UMMZ); 17 mi NW Custer, 3
102
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Table 19. — Geographic variation in coloration of the mid-dorsal pelage of adult Peromysctts maniculatus obtained in summer from the
Northern Great Plains.
|
Number and sex of specimens averaged |
Tone of color |
Percent red reflectance Percent green reflectance Percent blue reflectance |
|
33 (22 5,119 ) Mean |
P.m. 34.8 |
hiteus, Bennett County, South Dakota 48.2 27.8 24.0 |
|
SD |
±4.57 |
±0.30 ±0.15 ±0.19 |
|
55 (34,5,21?) Mean |
P. m. nehrascensis. Converse, Niobrara, and Weston counties, Wyoming 33.1 51.0 26.1 22.9 |
|
|
SD |
±5.15 |
±0.25 ±0.16 ±0.14 |
|
23 (13c?, 10 9 ) |
P. |
m. nehrascensis, Harding County, South Dakota |
|
Mean |
30.4 |
50.7 25.3 24.0 |
|
SD |
±2.93 |
±0.25 ±0.21 ±0.11 |
|
84(47 5,37 9) Mean |
28.2 |
P. m. nehrascensis. Black Hills, South Dakota and Wyoming 49.8 26.5 23.7 |
|
SD |
±4.58 |
±0.34 ±0.19 ±0.19 |
Table 20. — Color of dry samples of surface soil from the Black Hills (ar- ranged within counties in order of decreasing tone of color ) .
Locality
d) u
c -2 o^ o H o o
a; o
c B
<U O
S n3qq Ph (-1 >-,
3
ID O
3 05
SO
Ph be u
<D O
*^ s
3 B
O O
o o o
b 3 03
P-l ^ !=
Lawrence County
2 mi W Nemo 46.0
2 mi S Spearfish (A) 36.5
3 mi S Nemo 31.0
3 mi S Spearfish (B) 25.5
Timon Campgrounds 21.0
Meade County
3 mi SW Sturgis 39.5
Pennington County
Rapid City 97.5
3 mi SE Hill City 28.0
Bald Hills 27.5
Beaver Creek Valley 21.0
Custer County
Keystone — 50.0
Wind Cave National Park .... 32.0
Roby Springs 24.5
Custer 20.5
Flynn Creek 16.5
Fall River County
Hot Springs 36.0
1 mi SW Minnekahta 32.0
48.9 56.2 51.6 52.9 47.6
59.5
28.3 23.3 25.8 25.5 26.2
21.5
22.8 20.5 22.6 21.6 26.2
19.0
|
49.7 |
27.2 |
23.1 |
|
48.2 |
26.8 |
25.0 |
|
49.1 |
27.3 |
23.6 |
|
47.6 |
26.2 |
26.2 |
|
46.5 |
28.7 |
24.7 |
|
51.6 |
26.6 |
21.9 |
|
49.0 |
26.5 |
24.5 |
|
48.8 |
26.8 |
24.4 |
|
48.5 |
27.3 |
24.2 |
|
59.7 |
20.8 |
19.4 |
|
60.9 |
21.9 |
17.2 |
TURNER: MAMMALS OF THE BLACK HILLS
103
(UMMZ); 16 mi NW Custer, 1 (UMMZ); Harney Peak, 7240 ft, 4 (UMMZ); unspeci- fied locality, 9 (1 NRW, 3 SDMT, 5 UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill Cit>', 5300 ft, 12 (FMNH); 3.5 mi S, 0.5 mi W Ke\'stone, 5000 ft, 8; 5.75 mi N, 5.75 mi E Custer, 5220 ft, 19; Roby Springs, 4 mi N, 22 mi W Custer, 5400 ft, 2; Roby Canyon, 2 mi N, 22 mi W Custer, 5200 ft, 4; Squaw [Grace Coolidge] Creek, 14 (10 AMNH, 4 FMNH); Custer, 35 (32 AMNH, 3 FMNH); Harnev National Forest, 3-5 mi E Custer, 3 (UMMZ); 4 mi SW Custer, 9; Lightning Creek, 8 mi SW Custer, 1 (UMMZ); 4 mi NE Otis, 1 (UMMZ); 3 mi N Pringle, 5000 ft, 2 (UMMZ); Campbell's Ranch, Elk Mountain, 4800 ft, 14 (USNM); Wind Cave National Park, 10 (1 UW, 8 UMMZ, 1 WCNP); 6 mi N, 1 mi E Wind Cave, Wind Cave National Park, 4400 ft, 1; 5 mi N, 2 mi E Wind Cave, Wind Cax'e National Park, 4200 ft, 4; Buffalo Corral, Wind Cave National Park, 4300 ft, 8; 2.5 mi N, 5 mi E Wind Cave, Wind Cave National Park, 3700 ft, 5; Beaver Creek Can- yon, Wind Ca\'e National Park, 4200 ft, 4; Wind Cave, Wind Cave National Park, 4; Headquarters, Wind Ca\e National Park, 4100 ft, 8; Wind Cave Canyon, Wind Cave National Park, 4100 ft, 28; unspecified locality, 11 (UMMZ). Fall River Counti/: 1 mi N, 6 mi E Hot Springs, 3400 ft, 2; 1 mi N, 5.5 mi E Hot Springs, 3400 ft, 2; 5.5 mi E Minnekahta, 4050 ft, 4; 4 mi E Hot Springs, 3400 ft, 3; 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 72; 1.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 6; 1 mi S, 2 mi E Hot Springs, 3400 ft, 4; 2 mi S, 2 mi E Hot Springs, 3600 ft, 2; 4 mi S Hot Springs, 1 (UMMZ); 3 mi N, 2 mi E Edge- mont, 3500 ft, 1; 8 mi S Hot Springs, 3 (UXLMZ); Angostura Dam, 6.
WYOMLNG: Crook County: 6.5 mi SSE Alva, Bear Lodge Mountains, 2 (UMMZ); Devils Tower, flood plain Belle Fourche River, 33.50 ft, 2 (USNM); 15 mi ENE Sundance, 3825 ft, 11; Sand Creek, 1 (UW); 3 mi NW Sundance, 5900 ft, 12; 2 mi NW Sundance, 11; 1.5 mi NW Sundance, 5000 ft, 1; Sun- dance, 10 (USNM). Weston Counttj: 1.5 mi E Buck-horn, 6150 ft, 2; 4 mi E Four Corners, 2; 9 mi N, 1 mi E Newcastle, 5; 6 mi N New- castle, 1; 1 mi N Newcastle, 4800 ft, 11 (UMMZ); Newcastle, 12 (USNM).
Additional records.— SOUTH DAKOTA: Pennington Counttj: Rochford (BB); Medicine Mountain, 6200-6300 ft (Dice, 1942:3). WY- OMING: Weston County: Stockade Beaver Creek, 4300 ft (Dice, 1942:3).
Neotoma cinerea orolestes Merriam
Bushy-tailed Woodrat
Neotoma orolestes Merriam, 1894, Proc. Biol. Soc. Washington, 9:128, 2 July (type lo-
cality, Saguache Valley, 20 mi W Sagauche, Saguache Co., Colorado).
Neotoma cinerea orolestes — Goldman, 1910, N. Amer. Fauna, 31:104, 19 October.
Neotoma grangeri J. A. Allen, 1894, Bull. Amer. Mus. Nat. Hist., 6:324, 7 November (type locality, Black Hills, Custer, Custer Co., South Dakota).
Bu-shy-tailed woodiats are boreal mammals of higher altitudes and lati- tudes of the northern and western United States. In the Black Hills, Neotoma cin- erea occupies crevices in rocky outcrops, mine-shafts, and abandoned cabins from the boreal cap down through the transi- tion zone to about 4000 feet. These rats are active the year around and are com- mon throughout the study area. Vernon Bailey (1888:442) first noted this species in the Black Hills, indicating that its principal food seemed to be seeds of conifers and that large piles of gnawed cones often were scattered around en- trances to woodrat dens.
Woodrats are most abundant among ledges and vertical crevices in rocky outcrops that form the steep sides of valley and canyon systems; examples are Beaver Creek and Ditch Creek valleys, and Big and Little Spearfish canyons. Such areas support growths of ponderosa pine and creeping juniper. At lower ele- vations, where rocky terrain is less com- mon, buildings and other man-made fa- cilities often are inhabited. Mine shafts and caves also are frequented; I have observed N. cinerea in Wind Cave at depths of 325 feet below ground surface, and 2300 feet from any known entrance of the cave.
This species sometimes is active in late afternoon and one individual was taken by J. A. King at 16:30 hrs (MST) southeast of Hill City on 29 March 1946. This large cricetid builds nests that are globular or cup-shaped masses composed of dry grass, moss, leaves, and other fi- brous materials; these nests generally contain cones of pine and spruce, bones of various animals, and other debris. On 4 July 1967, I found a mandible of Canis lupus in a nest to the rear of Davenport Cave, and a subadult in Wind Cave
104
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
amassed numerous flashbulbs and can- dles within its nest.
Eight pregnant females taken in mid- June contained 3.6 (3-4) embryos that had an axerage crown-rump length of 32.1 (4-50). Three lactating females were captured in late July and two others were collected in August. A female ob- tained southwest of Sturgis on 14 August 1967 had six placental scars, and two others taken southwest of Lead in the same month had four placental scars each in the uterine walls. Testicular length of adult males varies seasonally. The a\'er- age testicular length of four males taken in June was 21 (16-30); whereas that of two captured in July was 12 each, and that of two obtained in August was 6 and 7, respectively. Juveniles are common in collections made from mid-June through August. Two suckling young, \\hich were only a few days old, were removed from the teats of a female taken in Dav- enport Cave on 26 June 1967; the young were nearly naked and darkly pigmented dorsally. Three suckling young ob- tained on 8 July from northeast of Deer- field were older and in fine, short juvenal pelage that was dark gray in color. Change from j^nenal to subadult pelage was evident in many specimens collected in late July, and several adults were un- dergoing annual molt o\'er the sides and rump in late July and early August. White, star-shaped spots were noted on the foreheads of many individuals, espe- cially on subadults.
Chambers (1948:8) observed a red squirrel kill a woodrat in the northern Hills near Lead. A female Neotoina from west of Nemo and a male taken southwest of Sturgis harbored ticks of the Ixodes ochotonae-augustus complex. Indi\'iduals from southwest of Lead were parasitized externally by two ticks, Ixodes spinipalpis Hadwen and Nuttall, and Dermacentor andersoni Stiles, two mites, Andwhelaps fahrenhohi (Ber- lese) and Haemogamasus amhulans (Thorell), two fleas, Orchopeas sexden- tatiis (Baker) and Monopsylus ivagneri
(Baker), and a chigger, Euschoengastia setosa ( Ewing ) .
Black Hills woodrats were described by J. A. Allen ( 1894a: 324-325) as a sepa- rate species, Neotoma grangeri, based on a series of 14 indi\iduals from Custer and two others from Glendale; 11 of these specimens were subadults. How- ever, in his revision of the genus Neo- tomcL Goldman (1910:105) indicated that topotypes of N. grangeri did not diff^er from typical N. c. orolestes, a con- clusion with which I concur.
Some secondary sexual variation is apparent among the Black Hills popula- tions of N. cinerea. Females are paler in tone of color and greater in breadth of interorbital constriction and length of maxillary tooth-row, but males are slightly larger in all other measurements. Because individuals are quite ^•ariable, the sexes were not treated separately in subsequent analysis.
Although no overall pattern of varia- tion in size is apparent, coloration of woodrats \'aries from place to place in the Hills. For example, specimens from Beaver Creek are paler in tone (29.5± 6.26), reflect reds (51.9±0.27) more in- tensely, and greens (23.7±0.13) and blues (24.2±0.20) less intensely than do specimens from Ditch Creek (24.7 ±3.96, 45.2±0.27, 28.1±0.15, and 26.5 ±0.29, respectively). Specimens from other localities represent gradations be- tween these extremes.
Neotoma cinerea orolestes differs from N. c. rupicoh on the plains to the south and cast in being darker and ha\- ing somewhat larger external and cranial dimensions. Specimens from the Black Hills arc on the average significantly darker in tone and reflect blues more in- tensely than do other representati\es of orolestes from west and nortli of the Hills (Fig. 14 and Table 21), but there were no significant differences in external or cranial measurements between these pop- ulations. Specimens from 0.5 mi E Buck- horn. Weston Co., Wyoming, and a sub- adult female from the Angostura Dam, Fall River Co., South Dakota, are some-
TURNER: MAMMALS OF TIIK BLACK HILLS
105
Fig. 14. Geographic variation in coloration of the mid-dorsal pelage of three groups of sum- mer-taken adult bushy-tailed \\oodrats. Neotoma cinerea orolestes averages much darker in tone than A^ c. rupicola, and specimens of N. c. orolestes from the Black Hills average much darker than non-Black Hills representatives to the west and north.
what more pale than typieal Black Hills representatives, but are still best referred to orolestes. Selected average external and cranial measurements of eight males and 15 females from the Black Hills are: total length, 359.1 ±20.67; tail length, 148.0± 10.31; hind foot length, 40.6 ± 2.12; ear length, 33.4±2.55; weight, 272.5 ±77.95; greatest length of skull, 48.3 ± 1.82; zygomatic breadth, 25.8±1.06; mas- toid breadth, 20.2 ±0.63; rostral length, 20.1 ±0.84; interorbital constriction, 5.9 ±0.28; length of incisive foramen, 11.2± 0.63; length of maxillary tooth-row, 9.9 ±0.49; depth of skull, 16.5 ±0.61.
Specimens examined (L34).— SOUTH DA- KOTA: Lawrence County: Little Spearfish Canyon, 2 mi S, L3 mi W Lead, 6000 ft, 1;
Little Spearfish Canyon, 2 mi S, H) mi W Lead,
0000 ft, 14; Little Spearfish Canyon, Savoy, 4 (USNM); Annie Creek, Black Hills, 6500 it, 1 (USNM); Deadwood, 2 (USNM); 2 mi W Nemo, 4700 ft, 2. Meade Counti/: Van- ocker Canyon, 3 mi S, 1 mi W Stiugis, 4600 ft, 1; Davenport Cave, 3 mi S, 0.5 mi W Sturgis, 4400 ft, 3. Pennington County: I3ark Canyon, 4 (AMNH); Diamond S Ranch, near Rapid City, 2 (UMMZ); Rapid City, 2 (1 NRW, 1 USNM); 15 mi SW Rapid City, 1 (NRW); 4.5 mi N, 2.5 mi E Deerfield, 4; Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 10; Ditch Creek, 14 mi W Hill Citv, 6400 ft. 10; 8 mi W Hill City, 7000 ft, 3 (UMMZ); Clendale, 2 (AMNH); Hill City,
1 (FMNH); Summit Peak, 3 mi SE Hill City, 5700 ft, 3 (UMMZ); Mushroom Rock, 4 mi SE Hill City, .5500 ft, 2 (UMMZ); Willow Creek, 4 mi SE Hill City, 5700 ft, 1 (UMMZ); unspecified locality, 1 (UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, 5300 ft, 1 (FMNH); Bull Springs, 2 mi N, 9 mi W Custer, 10 (UMMZ); Custer, 15 (2 USNxM, 12 AMNH, 1 FMNH); Housing Area, Wind Cave National Park, 4100 ft, 2; Wind Cave National Park, 7 (1 USNM, 1 WCNP, 5 UMMZ); Otis, 1 (UMMZ); Elk Mountain, 2 (USNM); unspecified locality, 7 (UMMZ). Fall River County: Angostura Dam, 1.
WYOMING: Crook County: Devils Tower,
2 (USNM); Sand Creek, 1 (USNM); 15 mi N Sundance, 1. Weston County: 0.5 mi E Buck- horn, 6100 ft, 5; 4 mi E Four Corners, 4; 3 mi N Newcastle, 1 (UW).
Clethrionomys gapperi brevicaudus
(Merriam)
Red-backed Vole
Evotomys gapperi brevicaudus Merriam, 1891, N. Amer. Fauna, 5:119, 30 July (type lo- cality, 3 mi N Custer, 6000 ft, Custer Co., South Dakota).
Clethrionomys gapperi brevicaudus — Bole and Moulthrop, 1942, Sci. Publ. Cleveland Mus. Nat. Hist., 5:153, 11 September.
The red-backed \ole is distributed in boreal and montane coniferous forest and its successional stages, and in brushy ri- parian habitats of North America. It inhabits the boreal cap of the Black Hills down to eknations of 4600 feet, but is most numerous above 5200 feet. It is active the year around. Although common in the Hills, no specimens are at hand from Meade or Fall Ri\'cr coun- ties.
Merriam named the subspecies, 1)revi-
106
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Table 2L — Geographic variation in coloration of the mid-dorsal pelage of adult Neotoma cinerea ol:)tained in summer from the
Northern Great Plains.
Number and sex
of specimens
averaged
H o o
o
c B
o a;
O
C O
ft 5b 2i
o
o a> 0;
OJ ,s
Ph^ ^
<a
|
23 (85,15$) |
N. c. orolestes. Black Hills, South |
|
Dakota and Wyoming |
|
|
Mean |
27.2 48.5 26.3 25.2 |
|
SD |
±4.31 ±0.36 ±0.24 ±0.23 |
|
21 (8(5,13$) |
N. c. orolestes, eastern half of Wyoming, |
|
and Harding County, South Dakota |
|
|
Mean |
37.0 49.6 27.0 23.4 |
|
SD |
±8.16 ±0.23 ±0.26 ±0.14 |
|
6(3c5,3$ ) |
N. c. rtipicola, western Nebraska |
|
Mean |
46.3 .50.8 26.4 22.8 |
|
SD |
±7.32 ±0.18 ±0.18 ±0.05 |
caudus, on the basis of tw^o specimens obtained near Custer on 18 and 21 July 1888. Bailey (1897:129) subsequently raised brevicaudtis to specific rank, be- cause: "Its range is isolated and widely separated from that of other members of the genus by open prairie country and a wide belt of Transition Zone." Bole and Moulthrop (1942:15.3) reestablished hrevicaudiis as a subspecies of C. gap- peri on the basis of additional material. As presently understood, l)revicaudus is restricted to the Black Hills.
Ecological sympatry of this vole with Microtus longicaudus and M. pennstjl- vanicus will be discussed later (see ac- counts of those species). Clefhrionomys primarily is an inhabitant of spruce, birch, and aspen woodlands, but also frequents drier pine-clad slopes. Indi- viduals generally are trapped near brush, woodpiles, boulders, fallen logs, or under low spreading branches of creeping juni- per. For example, Bailey trapped the holotype beneath a log in a deciduous thicket and captured a second individual adjacent to a log in pine timber on top of a ridge (Merriam, 1891:119). I cap- tured six voles in thickets of deciduous shrubbery along Spearfish Creek on 8 August 1967. Many additional speci-
mens were taken in brushy riparian habi- tats such as those found along Iron, Grizzly, Coldrun, Boxelder, Beaver, and Ditch creeks; soggy bog-Hke soils fre- quently were encountered in these areas.
In addition to moist spruce-filled can- yons and deciduous woodlands, red- backed voles also occupy rockslides, areas of secondary regeneration on burned over hillsides, or even areas of extreme exposure. In late November 1945, J. A. King trapped five Clefhrionomys in bare rocky areas among pine needles and other debris at the summits of Harney (7242 feet) and Summit (5700 feet) peaks. Occasionally, this species is active during the day. A male was taken about 10:00 hrs (MDT) west of Hill City on 12 June 1965, and another was captured at 19:. 30 hrs (MDT) in Beaver Creek Valley on 1 July 1965. Along with the meadow and long-tailed voles, Zaptis hndsonius, Sorex cinerens, Ferormjscus maniculaius, and Eutamias minimus are associates of red-backed \'oles.
Adequate reproductive data applica- ble to the Black Hills population of CJethrionomys are available only for July. However, abundance of subadults in mid-June and of jux enilcs in late Au- gust indicate that several litters are pro-
TURNER: MAMMALS OF THE BLACK HILLS
107
duced annually. A subadult female taken along Ditch Creek on 10 June 1965 car- ried three embr\os that had a crown- rump length of 4 each. Two adult fe- males obtained about this time showed no signs of reproductive actixity. The av- erage length of testes of nine males ob- tained in June was 9.0 (7-10); corre- sponding measurements of five subadults were 7.8 (6-9) . Eight of 15 adult females captured in July were pregnant; they had an average of 6.8 (5-9) embryos that were 10.1 (5-20) in crown-rump length. Testicular length of 12 males collected in July was 9.7 (7-11); that of three sub- adults was 8.3 (7-10). A lactating adult female was captured southwest of Rock- erville on 19 August 1965 and testicular length of six adult males obtained in August was 6.8 (6-8). Strangely, observa- tions concerning the occurrence of pla- cental scars in C g. hrevicaudus are lacking in field notes of collectors, even though notations of this type are entered for all other species; perhaps placental scars are more ephemeral in red-backed voles than in other small mammals.
Winter pelage is longer and softer than that of summer and the reddish brown dorsal stripe is much more dis-
tinct. Adults taken in mid-June and early July frequently were molting into the short grayish- to reddish-brown summer pelage; new hairs were enshrouded be- neath the overlying older pelage on both sides and dorsum. Because a distinctive moltline is not formed the process of pelage replacement is rather impercepti- ble. Fur of juveniles is more grayish in color than that of adults. Many sub- adults taken in summer were molting simultaneously over much of the body, whereas others had completed post- juvenal molt and were in fresh brownish pelage.
A male captured northeast of Custer on 19 June 1967 was infested with ticks, Dermacentor anclersoni Stiles.
Comparison of 27 male and 22 fe- male C. g. brevicaudus revealed no ap- parent secondary sexual dimorphism in coloration or in external and cranial di- mensions, nor was a pattern of variation in these characters evident in the Black Hills. Clethrionomys gapperi hrevicau- dus differs significantly from C. g. galei to the west in Wyoming in having a shorter tail and ear, longer hind feet, a higher skull, and darker pelage that reflects reds more intensely (Table 22).
Table 22. — Geographic variation in coloration of the mid-dorsal pelage of adult Clethrionomys gapperi obtained in summer from the
Northern Great Plains.
|
Number and sex of specimens averaged |
Tone of color Percent red reflectance Percent green reflectance Percent blue reflectance |
|
46 (24 5,22? ) |
C. g. brevicaudus. Black Hills, South |
|
Dakota and Wyoming |
|
|
Mean |
21.2 55.8 23.1 21.1 |
|
SD |
±2.19 ±0.34 ±0.29 ±0.19 |
|
12(65,69) |
C. a. loringi, Walsh County, North Dakota, |
|
Marshall County, South Dakota and Crow |
|
|
Wing County, Minnesota |
|
|
Mean |
21.0 57.4 22.3 20.3 |
|
SD |
±3.18 ±0.28 ±0.24 ±0.14 |
|
23 (16(5,79 ) |
C g. galei, Big Horn County, Wyoming |
|
Mean |
23.4 54.0 24.1 21.9 |
|
SD |
±2.25 ±0.26 ±0.19 ±0.15 |
108
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Clethrionomys gapperi hremcaudus also averages somewhat shorter in total length and greater in lengths of skull and rostrum, and in zygomatic and lamb- doidal breadths (Table 23). From C. g. loringi to the north and east, brevicaiiclus differs significantly in longer hind feet, greater lambdoidal breadth and a higher skull; it is also smaller in total length and length of tail, and greater in lengths of ear, skull, and rostrum and in zygo- matic breadth. Measurements additional to those given in table 23 for 49 adult red-backed voles from the Black Hills are: weight, 25.4±;4.48; interorbital con- striction, 3.9 ±0.13; length of incisive foramen, 4.9 ±0.26; length of maxillary tooth-row, 5.4 ±0.25.
Specimens examined (229).— SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- yon, 6 mi S, 2 mi W Spearfish, 4600 ft, 6; Tinton, 5900 ft, 1; 2 mi S Tinton, 6100 ft, 22; Deadwood, 1 (USNM); Dumont, 6100 ft, 5 (USNM); 2 mi W Nemo, 4700 ft, 4; Nemo, 4700 ft, 1. Pennington County: Jim Creek, east of Merritt, 1 (NRW); Rapid" Creek, 1.5 mi W Rochford, 1 (UMMZ); Diamond S Ranch, near Rapid City, 1 (UMMZ); Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 21; 3 mi N, 7 mi W Deerfield, 6900 ft, 1; 1 mi S, 2 mi E Deerfield, 6400 ft, 1; McVey Burn, 1 (NRW); 20 mi N Elk Mountain, 6000 ft, 2 (USNM); Ditch Creek, 14 mi W Hill City, 6400 ft, 3; Hill Citv, 1 (AMNH); Palmer
i mi SE Hill mi E Sylvan mi S, 0.5 mi N, 5.75 mi E
Gulch, 3 mi SE Hill City, 5300-.5400 ft, 11 (UMMZ); Willow Creek, Nelson's Place, 4 mi SE Hill City, 5300-5400 ft, 13 (UMMZ); Harney Peak, 5 mi SE Hill City, 7240 ft, 1 (UMMZ); Rockerville, 1 (UMMZ); 3 mi S,
1 mi W Rockerville, 3; Keystone, 1 (NRW); 17 mi NW Custer, 11 (UMMZ); 16 mi NW Custer, 22 (UNLMZ); 16 mi SW Rapid Citv, 1 (UMMZ); unspecified locality, 2 (UMNIZ). Custer County: Palmer Gulch, City, 5300 ft, 9 (FMNH); 0.5 Lake, 62.50 ft, 1 (UMMZ); 3.5 W Keystone, 5000 ft, 1; 5.75 mi Custer, 5220 ft, 21; 3 mi N Custer, 6000 ft,
2 (USNM); Custer, 21 (18 AMNH, 3 FMNH); Bull Springs, 2 mi N, 9 mi W Custer, 7 (UMMZ).
WYOMING: Crook County: 3 mi NW Sundance, 5900 ft, 3; Rattlesnake Creek, 3 (USNM). Weston County: 1.5 mi E Buck- horn, 6100 ft, 21; 12 mi SE Newcastle, 1 (UMMZ).
Microtus longicaudus longicaudus
(Merriam)
Long -TAILED Vole
Arvicola (Mijnomes) longicaudus Merriam, 1888, Amer. Nat., 22:934, October (type locality, Custer, 5500 ft, Custer Co., South Dakota ) .
Microtus (Mynomes) longicaudus — T- A. Allen, 1895, Bull. Amer. Mus. Nat. Hist., 7:266, 21 August.
The long-tailed \'ole occurs on the boreal cap of the Black Hills and inhab-
Table 2.3. — Geographic variation in selected external and cranial measurements of adult Clefliriono-
mys gapperi from the Northern Great Plains.
|
Number and sex of specimens |
r" |
*-i |
Ifoot th th itest th nil )matie dth ral th bdoidal dth |
|
|
iS w^ |
-~ tc |
c « ^ S^' a3 ^'^ a o ^ S^ s « |
.— o |
|
|
averaged |
e2^ |
H^ |
XM w^ O^'t ^^ ^J J^ |
on T3 |
|
49(27,^,22 9 ) |
G.g |
. brevicaudus. Black Hills, South Dakota and Wyoming |
||
|
Mean |
134.0 |
34.5 |
18.7 14.9 24.5 13.6 8.5 11.6 |
9.6 |
|
SD |
±8.60 |
±3.37 |
±0.95 ±1.63 ±0.76 ±1.49 ±0.41 ±0.36 |
±0.26 |
|
19(1U,8$) |
G.g. |
loringi, Walsh Coxmty, North Dakota, Marshall County, Sotith Dakota and Crow Wing County, Minnesota |
||
|
Mean |
1.35.2 |
35.8 |
18.2 14.1 23.7 13.2 8.3 11.3 |
9.1 |
|
SD |
±9.36 |
±3.69 |
±1.28 ±1.13 ±0.71 ±0.64 ±0.33 ±0.22 |
±0.22 |
|
23(16^,7$ ) |
G. g. galei, Big Horn County, Wyoming |
|||
|
Mean |
140.7 |
38.8 |
18.3 16.4 24.3 13.3 8.3 11.5 |
9.3 |
|
SD |
±8.83 |
±3.72 |
±0.88 ±1.26 ±0.43 ±0.32 ±0.21 ±0.19 |
±0.24 |
TURNER: MAMMALS OF THE BLACK HILLS
109
its riparian environs bordering cold streams that descend into the lower semi- arid transition zone. It is widely dis- tributed but none has been taken as yet in Fall Ri\er County. Although speci- mens have been obtained in the Hills as low as 4200 feet, M. longicaudus is most abundant at elevations above 6000 feet. The Black Hills population is iso- lated from other generally montane pop- ulations westward and southward in Wy- oming. The species does not hibernate and indi\'iduals have been captured in all seasons.
A distinct pattern of ecological sepa- ration between M. longicaudus and M. pennsyJvanicus is not readily discernible in the Hills. W. W. Granger wrote about them as follows, "I found these mice [A/. pen}isyJvanicus] in the same locali- ties as the other species [M. longicau- dus]. Some were caught on a hillside which was covered with aspens, and the rest along the banks of a creek" (J. A. Allen, 1895a: 267). Both species fre- quently are captured in the same trap- line in the mesic Northern Hills. For example, a Sherman live-trap placed be- neath the overhanging bank of Little Spearfish Creek in the Timon Camp- ground obtained one M. longicaudus and one M. pennsylvanicus within an hour ( 19:30-20:30 hrs MDT) on 31 July 1968. Vegetation in the vicinity of the trap was composed of mosses, grasses, and deciduous shrubs.
At higher elevations, M. longicaudus ranges farther from riparian situations than does M. pennshjvanicus, which in- habits moist meadows and edges of spruce and aspen forest. In the latter habitat, long-tailed \'oles overlap Cleth- lionomys gapperi. At lower elevations, M. longicaudus is more restricted to streamside environments and lowland marshes of rushes, sedges, and cattails, or other moist areas of much brush and fallen logs. In these areas, M. pennsyl- vanicus ranges farther from riparian situ- ations and is more tolerant of the drier meadows. In addition to M. pennsylvani- cus and C. gapperi, other species trapped
in association with long-tailed voles are Peroniyscus nianiculatus, Eutamias mini- mus, Zapus hudsonius, and Sorex cine- reus.
Reproducti\e data for long-tailed voles in the Black Hills are adecjuate only for July. Of two females taken west of Hill City in mid-June 1965, one carried five embryos which were 10 in crown- rump length; the other showed no signs of reproductive activity. Testicular length of a male obtained on 28 June 1967 was 12. Fifteen of 23 females cap- tured in July were pregnant; there was an average of 4.7 (4-6) embryos in each, which were 10.3 (3-24) in crown-rump length. One female collected northwest of Deerfield on 8 July 1965 had eight placental scars; another taken east of Buckhorn on 18 July 1951 was lactating. An additional female trapped southwest of Lead on 30 July 1968 had a swollen uterus, possibly indicating recent im- plantation. Testes of 12 adult males cap- tured in July had an average length of 11.2 (10-12). On 9 August 1967, two females were captured south of Spear- fish; one carried six embryos that were 4 in crown-rump length, whereas the other was lactating and had seven pla- cental scars. Juvenile and subadult long- tailed voles are common in collections made throughout July and August, and many of the pregnant females were sub- adults.
Five young voles taken in Beaver Creek Valley in early July were in Juv- enal pelage, which was fine, soft, and dark. Twelve juveniles collected through- out July either had completed post- juvenal molt, or replacement of juvenal pelage was in process with new hairs exposed beneath the old pelage on the sides and dorsum. In the same period, three subadults were molting to an adult pelage. In adults, change from old, faded winter pelage to fresh summer pelage was noted in late June and early July; irregular patches of new hair oc- curred on the rump and over the shoul- ders.
A male trapped in Beaver Creek
110
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Valley on 8 July 1965 harbored ticks of the Ixodes ochotonae-angustus complex.
Significant secondary sexual dimor- phism was not evident in comparison of 14 adult males with 17 adult females from the Black Hills, although males are slightly larger in most measurements and females (50.9 ±0.25) reflect reds more intensely than males (48.8±0.15). A distinct pattern of variation in colora- tion or size within the Hills populations is not discernible.
Bailey (1900:48) noted that M. /. longicaudus and M. I. mordax are "very similar" in external and cranial dimen- sions. After examining specimens from Idaho and throughout Wyoming, Long (1965:655) placed mordax in the synon- omy of longicaudus, pointing out that C. H. Merriam selected an atypically dark vole with long ears as the holotype. When 31 long-tailed voles from the Hills were compared with 29 from north- central Wyoming (Big Horn, Fremont, Johnson, Sheridan, and Washakie coun- ties ) and 80 from southeastern Wyoming (Albany, Carbon, Converse, Laramie, and Natrona counties); the former aver- aged darker in tone and possessed longer ears and maxillary tooth-rows (Table 24).
Even though the pelage of M. longi- caudus is extremely uniform in reflect- ance of various hues, this species is quite variable throughout its range in all other parameters. For example, relative to the Black Hills population, 28 individuals from Carbon County, Wyoming, are darker in tone of color (21.0±1.98), 17 specimens from Natrona County have a similar average length of ear ( 15.8± 1.59), and single representatives from Washakie and Sheridan counties each have longer maxillary tooth-rows (7.5 and 7.0, respectively). Thus, the above- mentioned apparent differences attribut- able to the Black Hills population of M. longicaudus easily fall within the range of variation of populations southward and westward of the Hills, and I must concur with the findings of Long {loc. cit. ) . Average color and cranial measure-
ments additional to those given in Table 24 for 31 adults from the Black Hills are: percent red reflectance, 50.0±0.23; percent green reflectance, 26.7 ±0.18; percent blue reflectance, 23.3±0.16; weight, 41.9±7.04; interorbital constric- tion, 3.8±0.15; length of incisive fora- men, 5.0±0.23.
Specimens examined (172). — SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- yon, 3 mi S Spearfish, 4200 ft, 2; Tinton, 5900 ft, 1; 2 mi S Tinton, 6100 ft, 4; Little Spear- fish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 3; Cheyenne Crossing, 8 mi SW Lead, 1 (SDSU); Boxelder Creek, Steamboat Rock Campgrounds 1 (NRW); Dumont, 1 (USNM). Meade County: 7 mi S, 1 mi W Sturgis, 4700 ft, 1. Pennington County: Rapid Creek, 1.5 mi W Rochford, 4 (UMMZ); Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 17; Ditch Creek, 14 iTii W Hill City, 6400 ft, 2; Redfern, 1 (USNM); 20 mi N Elk Mountain, 6000 ft, 12; 17 mi NW Custer, 7 (UMMZ); 16 mi NW Custer, 23 (UMMZ); Palmer Gulch,
3 mi SE Hill City, 5300-5400 ft, 7 (UMMZ); Nelson's Place, 4 mi SE Hill City, 5300-5400 ft, 2 (UMMZ); Palmer Gulch, 5 mi SE Hill City, 2 (UMMZ); unspecified locality, 5 (UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, 5300 ft, 4 (FMNH); 16 mi W Custer, 7 (USNM); Custer, 6 (2 USNM, 4 AMNH); 4 mi E Custer, Harney National Forest, 1 (UMMZ); Elk Mountain, 4900 ft,
4 (USNM); 4 mi SW Custer, 1; 18 mi SW Custer, 5300 ft, 5 (USNM); Bull Springs, 2 mi N, 9 mi W Custer, 8 (UMMZ).
WYOMING: Crook County: Bear Lodge Mountains, Warren Peak, 6000 ft, 1 (USNM); Bear Lodge Mountains, 6.5 mi SSE Alva, 1 (UMMZ); 3 mi NW Sundance, 5900 ft, 7; 1.5 mi NW Sundance, 5000 ft, 1; Sundance, 6000 ft, 12 (USNM); Rattlesnake Creek, 6000 ft, 2 (USNM). Weston County: 1.5 mi E Buckhorn, 6150 ft, 15; 4 mi E Four Corners, 1.
Additional record. —SOUTH DAKOTA: Pennington County: Rochford (BB).
Microtus ochrogaster haydenii (Baird) Prairie Vole
Arvicola (Pedomys) haydenii Baird, 1858, in Reports of explorations and surveys . . . from the Mississippi River to the Pacific Ocean . . ., 8(1):543, 14 July (type lo- cality, Fort Pierre, Stanley Co., South Da- kota).
Microtus ochrogaster haydenii — Osgood, 1907, Proc. Biol. Soc. Washington, 20:48, 18 April.
Microtus ochrogaster, in contrast to
TURNER: MAMMALS OF THE BLACK HILLS
111
C3
O
o
0) J5
C O
CO
S O
"Sb c o
CO
03
C3
'c 3
^3
C
c
o
c o
XI
•n
>
:a &
5c o
O
<;
H
|
tpd.^p |
lo CO |
CD CO CO |
in ^ CO |
|
|
d d |
d d |
d d |
||
|
u"ns |
^ +1 |
^ +1 |
^ +1 |
|
|
A\o.i-q}ooj |
00 M |
cq 1^ CO |
02 CD (M |
|
|
X.IB[[IXUIU |
d d |
d d |
CD d |
|
|
JO ijiSuaq |
+1 |
+1 |
+1 |
|
|
CQ |
00 |
l> |
||
|
mSuar |
t^ TP |
00 CO |
CD CO |
|
|
00 d |
00 d |
00 d |
||
|
[B-Hsoy |
+1 |
+1 |
+ 1 |
|
|
1— ( |
CD |
CD |
||
|
ipptJOKj |
^. ^ |
in in |
cq Ti^ |
|
|
oi d |
(>i d |
c4 d |
||
|
piOpC[lUB'^ |
^ +1 bC c |
^ +1 |
^ +1 |
|
|
o in |
i-H |
05 |
||
|
ippea-iq |
dWv 15.6 ±2.0 |
(M 00 |
o m |
|
|
OpV!UT08X2 |
.5^ +1 |
c -^ +1 |
||
|
a |
s o |
S o |
||
|
4-t o |
^ |
^ |
||
|
I[n>is JO |
rt CO 00 |
'rt |
in CD 05 |
|
|
i[43u3[ |
Qt^d |
^-' |
i> d |
53 1^ o |
|
}S9;Ba.io |
c a.) o |
oi II |
||
|
o |
^ |
|||
|
CA) |
t |
•*-t |
||
|
c/f |
o |
o |
||
|
ipSu9[ |
Hill 5.8 1.48 |
:z, |
o |
|
|
.iBa |
tp' ^ |
'^ 1-H |
||
|
^ +1 |
-" II |
|||
|
^ n |
||||
|
o |
in |
o |
||
|
i{j§ua[ |
CO 00 -(■ d |
CO I> ^ d |
CD cq d ^' |
|
|
jooj puiH |
oi II |
oi II |
oi II |
|
|
in |
00 |
t- |
||
|
ipguaj |
CO ^ |
Tf< OQ |
(M O |
|
|
II«X |
d ^ |
t^ d |
ai in |
|
|
in +1 |
in II |
in II |
||
|
t- |
C<I |
a> |
||
|
iflSugj |
a CT) |
CD O |
in CO |
|
|
00 00 |
d 00 |
d 00 |
||
|
F^ox |
^+1 |
^+1 |
^+1 |
|
|
cq |
^ |
I-H |
||
|
.lOIOO 10 |
O 00 |
.-1 05 |
1> 1>; |
|
|
'-^ ci |
in oi |
c4 <N |
||
|
auojL |
oi II |
<N II |
(M II |
X
C C aj
c8 .5 '^
"^ ^ 2
<u a^ 53
a '^ ^
3 O
o
I-H
CO
Q
o
2S
Q
05 CM
Of
?— I
o
00
Q en
112
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
other members of the genus that occur in the study area, is relatively tolerant of xeric conditions and is widely distrib- uted wherever suitable grassy habitats are available on the Great Plains of cen- tral North America. Prairie \'oles have been taken in all counties within the Black Hills, and are found in the drier valley systems that penetrate the foothill transition zone. They occur up to 5000 feet, but are most abundant below 4400 feet along the interface of the surround- ing prairie. Wasteland, dry ditches along roadways, and lightly grazed pastures also are inhabited by this vole. Vernon Bailey (1888:445) reported the first specimens of M. ochrogaster in the Black Hills region.
From 7 to 11 August 1967, a survey was taken along the floor of Big Spearfish Canyon, out onto the adjacent prairie. The vegetation of this moist northern canyon system is rather diverse. At higher elevations, ponderosa pine is fairly restricted to the arid bluffs and white spruce dominates the mesic can- yon bottoms, along with rushes, sedges, dense stands of tall grasses, deciduous shrubs, and quaking aspen; Clethriono- rnys gapped, Microtus longicauclus, and M. pennsijlvanicus were taken in these areas. Descending along the canyon, conditions gradually became less mesic. Deciduous elements such as bur oak, willow, paper birch, quaking aspen, box- elder, bunchberry, and alder are abun- dant. The understory of parklike wood- lands and small meadows is composed of wild gooseberry (Rihes missouriense) , wild strawberry (Fragaria virginiana), chokecherry, poison ivy, Solomon's seal, starry solomonplume {Smilacina stel- lata), yellow lady's slipper (Cijpripe- (Uiim sp. ), white geranium (Geranium bicknellii), black-eyed susan, sunflower, burdock (Arctium minus), cinquefoil (Potentilla sp.), and meadow sweet ( Spiraea lucida ) . Along Spearfish Creek, and adjacent moist areas at these inter- mediate elevations are snowberry, horse- tail (Equisetum sp.) flowering fern (Os- munda sp.), wood fern (Dryopteris sp.).
rushes, and sedges; Microtus longicau- dus and M. pennsylvanicus were ob- tained in these areas, but Cleithriono- mys no longer was taken. Below 4000 feet, upward extensions of the xeric prairie were encountered as e\ idenced by the presence of thistles, soapweed, mustards, and an occasional pricklypear cactus. Prairie voles were taken under these conditions, whereas the other three microtines were no longer encountered.
Fifty-eight individuals were captured southwest of Minnekahta ( 13-18 June 1967) on slopes that form the southern limits of the Black Hills. The area con- sists of arid grassy pastureland, expanses of dryland farming (mostly small grains), and a series of ridges upon which ponderosa pine and western red cedar grow. Most of the voles were taken along fencerows bordering agri- cultural fields, in vegetation composed of bluegrass, brome grass, cord grass, west- ern wheat grass, and scattered forbs. Vole runways were well-defined but lo- calized, and piles of grass cuttings were common. Specimens collected near Hot Springs were on grass ridges and in ap- ple orchards that flank the Fall River.
Wind Cave National Park extends through the gently rolling foothills of the transition zone, grading into the more rugged topograph)' of the southeastern portion of the Black Hills. Here, where Al. ochrogaster is one of the commonest small mammals, the mixed-grass prairie of the plains intermingles with the pon- derosa pine of the higher elevations. Chokecherry, wild plum (Prunus ameri- cana), wild rose, hackberry, and sage- brush are interspersed with prairie grasses and forbs. Other mammals trapped in the runways of this species were Pcromyscus maniculatus, Perogna- thus hispidus, Mus musculus, and Reith- rodontomys megalotis.
Reproductive activity was substantial from late spring throughout the summer, and continued at least into early autumn. Three females captured in late March and early April carried 4.3 (4-5) em- bryos that had an average crown-rump
TURNER: MAMMALS OF THE BLACK HILLS
113
length of 5.7 (2-10). Reproducti\ c> data are unavailable for May, early June, and July (except for one female captured on 26 July 1968 that carried three embryos which were 8 in crown-rump length). Field work dining these periods \\'as un- dertaken at higher ele\ations where prairie voles do not occur, thus explain- ing the lack of data. Sixteen of 26 fe- males taken in the last half of June were pregnant, containing 3.5 (1-6) embryos that had an a\ erage crown-rump length of 10.5 (3-20). Two other females had 3.0 (2-4) placental scars, and another female had an enlarged uterus, which possibly indicates recent implantation. A\erage testicular length of 22 adult males obtained in this period was 11.0 (7-17). In the latter part of August, three of nine females examined contained fetuses. The average number of fetuses was 3.7 (3-4); the average crown-rump length of the fetuses was 11.3 (6-20). Two other females had five placental scars each. The average length of the testes of 11 males obtained in August was 13.0 (10-18). Five of eight females cap- tured in early September were pregnant, and contained an average of 3.8 (3-5) embryos that were 15.2 (8-22) in crown- rump length; four placental scars were manifest in the reproductive tract of one other female. The average testicular length of five adult males obtained about this" time was 12.6 (9-17). Juveniles and subadults are common in collections made from April through September; several of the gravid females were nearly mature subadults.
Due to the high reproductive rate of this species, seasonal aspects of molt were difficult to discern. Juveniles were in a fine dark pelage, although many were commencing the post-juvenal molt. As voles attained about three-quarters of adult size and adult pelage replaced that of the subadults, new hairs could be observed underlying the older fur on the dorsum and sides, and occasionally on the venter. Irregular patches of molt were noted on adults in all months from D which specimens were examined. Pelage
replacement in adulls s(^ems to be more erratic than in subadults and possibly extends over a greater period of time.
Prairie voles from the Hills are heav- ily infested \\ilh various types of ecto- parasites as follow: a louse, Iloplopleiira acanthopus (Burmeister); a tick, Ixodes spinipaipis Hadwen and Nuttall; two fleas, Orchopeas leucopus (Baker), and MonopsyUus ica^neri (Baker); two chig- gers, Euiromhicxda alfreddugesi (Oude- mans), and Euschoengastia setosa (Ew- ing); and seven mites, Dermacams hy- pudaei (Koch), Mycoptes musculinus (Koch), Haemolaelaps jahrenhohi Ber- lese, Androlaelaps jahrenhohi (Berlese), Laelaps microti (Ewing), Ornithonyssus hacoti (Hirst), and Haemogamasus am- hulans (Thorell).
Examination of 26 adult male and 23 adult female M. ochrogaster from the Black Hills revealed no secondary sexual differences in coloration, external meas- urements or most cranial dimensions. However, females (6.9±0.32) are sig- nificantly greater than males (6.6d=0.29) in length of maxillary tooth-row. A pat- tern of variation within the Hills was not evident. When comparisons were made with populations from other areas in South Dakota and Wyoming, the great variabifity of M. ochrogaster became ap- parent (Tables 25 and 26). Characters seem to vary independently and no def- inite trends are discernible, although specimens from the Hills tend to be darker in tone of color and relatively large in external and cranial dimensions. Measurements additional to those given in Table 26 for 44 adult prairie voles from the Hills are: hind foot length, 20.2±0.94; lambdoidal breadth, 12.4db 0.39; interorbital constriction, 4.0 ±0.13; length of incisive foramen, 5.0+0.32; and length of maxillary tooth-row, 6.7 it 0.34.
Specimens examined (214).— SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- yon, 1.5 mi S, 0.5 mi E Spearfish, 3800 ft, 6. Meade County: Black Hawk, 22 (AMNH). Pennington County: 8 mi NE Rapid City, 4 (UMMZ); 4 mi N Rapid City, 2 (AMNH); South Canyon, 3 mi W Rapid City, 13 (6 AMNH, 7 NRW); Rapid City, 9 (4 USNM,
114
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Table 25. — Geograpliic variation in coloration of the mid-dorsal pelage of adult Microtus ochrogaster obtained in summer from the
Northern Great Plains.
|
<0 (D |
OJ |
||
|
O O |
o |
||
|
Number and sex |
|||
|
c < |
01 k- |
0) o 0; C CJ |
<D O |
|
or specimens averaged |
c o H c o |
Perc red refle Perc gree refle |
Perc blue refle |
|
41 (22 5,19$) |
Black Hills, South Dakota and Wyoming |
||
|
Mean |
20.0 |
48.0 27.5 |
24.5 |
|
SD |
±2.97 |
±0.20 ±0.18 |
±0.16 |
|
20(115,95) |
Harding County, South Dakota |
||
|
Mean |
21.8 |
48.1 27.3 |
24.6 |
|
SD |
±3.00 |
±0.26 ±0.18 |
±0.15 |
|
8(45,45) |
Bennett County, South Dakota |
||
|
Mean |
23.6 |
47.6 26.8 |
25.6 |
|
SD |
±4.72 |
±0.28 ±0.21 |
±0.13 |
|
4(25,25) |
Campbell Countv, Wyoming |
||
|
Mean |
19.9 |
52.7 24.6 |
22.7 |
|
SD |
±1.84 |
±0.24 ±0.13 |
±0.20 |
|
2(15,15) |
Weston County, Wvoming |
||
|
Mean |
25.5 |
46.1 28.5 |
25.4 |
|
SD |
±2.12 |
±0.03 ±0.10 |
±0.06 |
|
5(45,15) |
Converse County, Wyoming |
||
|
Mean |
21.2 |
48.1 27.4 |
24.5 |
|
SD |
±2.41 |
±0.42 ±0.38 |
±0.08 |
|
16 (85,85) |
Niobrara Countv, Wyoming |
||
|
Mean |
23.2 |
51.3 25.7 |
23.0 |
|
SD |
±3.12 |
±0.33 ±0.23 |
±0.16 |
5 NRW); 1 mi SW Rapid City, 2 (UMMZ); West Dark Canyon, 1 mi S, 4 mi W Rapid City, 2 (AMNH); near Rim Rock, 2 (AMNH); Bald Hills, south Pactola, 4 (USNM); Spring Creek, 4 mi ENE Rockerville, 3600 ft, 7 (UMMZ); Spring Creek, south Rapid City, 10 (8 AMNH, 2 MHM); unspecified locality, 1 (SDMT). Custer County: Beaver Creek, 3 mi NW Wind Cave National Park, 1 (UW); Wind Cave National Park, 9 (7 UMMZ, 2 WCNP);
6 mi N, 1 mi E Wind Cave, Wind Cave Na- tional Park, 4400 ft, 1; 2.5 mi N, 5 mi E Wind Cave, Wind Cave National Park, 3700 ft, 5; 1.5 mi N, 1.5 mi W Wind Cave, Wind Cave National Park, 3250 ft, 4; Buffalo Corral, Wind Cave National Yark, 4300 ft, 1; Headquarters, Wind Cave National Park, 4100 ft, 8; Housing Area, Wind Cave National Park, 4100 ft, 2; Wind Cave Canyon, Wind Cave National Park, 4100 ft, 3; Elk Mountain, 4800 ft, 3 (USNM); unspecified locality, 9 (UMMZ). Fall River County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 58; 1 mi N, 5.5 mi E Hot Springs, 3400 ft, 4; 1 mi N, 6 mi E Hot Springs, 3400 ft, 11; 4 mi E Hot Springs, 3400 ft, 3; Hot Springs, 1 (UW); 2 mi S, 2 mi E Hot Springs, 3600 ft,l.
WYOMING: Crook County: 15 mi NE Sundance, 3825 ft, 1; 1.5 mi NW Sundance, 5000 ft, 1; Sundance, 2 (USNM); Sand Creek, 3750 ft, 1 (USNM). Weston County: 1 mi N Newcastle, 4800 ft, 1 (UMMZ); Newcastle, 1 (USNM).
Additional reco/f/.— SOUTH DAKOTA: Lawrence County: Spearfish (BB).
Microtus pennsylvanicus insperatus (J. A. Allen)
Meadow Vole
Arvicola insperatus J. A. Allen, 1894, Bull. Amer. Mus. Nat. Hist., 6:347, 7 December ( type locality, Custer, Custer Co., South Dakota).
Microtus pennsylvanicus insperatus — Anderson, 1943, Canadian Field-Nat., 57:92, 17 Oc- tober.
The meadow vole occupies streaniside habitats and mesic areas adjacent to reservoirs and other impoundments throughout the Northern Great Plains,
TURNER: MAMMALS OF THE BLACK HILLS
115
Table 26. — Geographic variation in selected external and cranial measurements of
Microtiis ochrogaster from the Northern Great Plains.
adult
|
Nimiber and sex of specimens averaged |
Total length |
Tail Length |
Ear length Weight Greatest length of skull Zygomatic breadth |
Rostral length |
Skull depth |
|
44 (24 5,20 9) |
Black Hills, South Dakota and Wyoming |
||||
|
Mean |
158.1 |
37.5 |
13.7 48.5 27.9 16.1 |
9.0 |
11.0 |
|
SD |
±9.86 |
±3.47 |
±1.66 ±6.37 ±1.06 ±0.69 |
±0.51 |
±0.34 |
|
20(11^,9?) |
Harding County, South Dakota |
||||
|
Mean |
158.8 |
36.2 |
13.1 46.7 27.9 15.7 |
8.9 |
10.9 |
|
SD |
±9.39 |
±4.19 |
±1.98 ±4.68 ±1.06 ±0.67 |
±0.42 |
±0.32 |
|
8(45,49) |
Bennett County, South Dakota |
||||
|
Mean |
152.2 |
34.2 |
11.7 36.0 27.2 15.1 |
8.8 |
10.7 |
|
SD |
±8.38 |
±4.83 |
±1.03 ±2.88 ±0.88 ±0.39 |
±0.26 |
±0.47 |
|
4 (25,29) |
Campbell County, Wyoming |
||||
|
Mean |
145.7 |
34.7 |
14.2 37.2 26.8 15.2 |
8.2 |
10.5 |
|
SD |
±3.86 |
±4.99 |
±2.06 ±4.03 ±0.82 ±0.47 |
±0.37 |
±0.41 |
|
2 (15,19) |
Weston County, Wyoming |
||||
|
Mean |
171.5 |
41.0 |
13.0 49.5 28.8 16.3 |
9.3 |
11.3 |
|
SD |
±7.78 |
±0.00 |
±0.00 ±7.99 ±1.20 ±1.38 |
±0.71 |
±0.32 |
|
5(45,19) |
Converse County, Wyoming |
||||
|
Mean |
160.8 |
40.2 |
14.8 47.7 28.3 15.8 |
8.9 |
11.0 |
|
SD |
±4.21 |
±4.27 |
±1.64 ±3.59 ±1.01 ±0.64 |
±0.44 |
±0.24 |
|
16(85,89) |
Niobrara County, Wyoming |
||||
|
Mean |
167.1 |
39.7 |
14.1 47.3 28.1 16.2 |
9.0 |
11.1 |
|
SD |
±9.71 |
±4.57 |
±1.54 ±6.74 ±1.27 ±0.89 |
±0.47 |
±0.50 |
Altitudinal range within the Black Hills is from 3700 to 6500 feet, but the species is most common above 5000 feet. Range of tolerance of environmental conditions seems to be greater in Microtus pennsyl- vanicus than in any other microtine from the study area. Meadow voles occur from the mesic boreal cap of the Hills, down through the semiarid foothill tran- sition zone, and onto the arid plains in suitable riparian habitats; they are active in all seasons. Although this microtine is the commonest member of the genus in the Black Hills, specimens are lacking from Meade and Fall River counties.
Streamside habitats in canyons of spruce, aspen, willows, sedges, grasses, rushes, deciduous bushes and scattered forbs are favored by this species at higher elevations; 52 meadow voles ( 19 males and 33 females) were taken south of Tinton under these general conditions
in mid-July 1961. Swampy lowlands, borders of beaver ponds and cattail ( Typha sp. ) marshes, and moist mea- dows of bluegrass, spearmint {Mentha spicata), stinging nettle (Urtica sp.), red clover {Trifolium pratense), and as- sorted forbs are occupied at lower ele- vations. Individuals captured northeast of Sundance (early July 1947), west of Nemo (late June 1947) and in Wind Cave National Park (summer 1968) are from such environs. Traps placed among mossy rocks and ferns under overhanging stream banks tliroughout the Hills also proved to be productive in capturing meadow voles.
Due to its wide ecological distribu- tion, Microtus pennsylvanicus is syntopic with three other species of voles in the Black Hills region. Association of this species with Microtus longicaudus al- ready has been discussed (see account
116
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
of that species). Morris (1969:299) re- ported an almost complete habitat sepa- ration between M. penmijlvanicus and Clethrionomys gapperi in Saskatchewan, with the former being confined to grass- land and the latter to aspen woodland. In the Hills, however, both species fre- quently were taken in the same habitat. Although M. pennsylvanicus was not captured in stands of aspen or other woodlands, C. gapperi often was ob- tained in grassland and riparian situa- tions where brush, woodpiles, and fallen logs were common; here, the two species occurred side by side. At lower eleva- tions, where the meadow vole and prairie vole occur together. M. penmijlvanicus inhabits the lower, wetter areas and M. ochrogaster the drier places. For ex- ample, in Wind Cave National Park, the meadow vole occupies the immediate brushy banks of Beaver Creek, whereas the prairie vole occupies the gi-assy up- land adjacent to the creek. In addition to the above named species, Peromijscus maniculatiis, Eiitamias minimus, Zaptis litidsoniiis, and Sorex cinereus were taken in the same ti'aplines with M. pennsyl- vanicus.
Reproductive data for meadow voles in the Black Hills are available only for the period from mid-June through early August. Bailey (1924:528) indicated that parturient female M. pennsylvaniciis have been taken in all seasons. Four of six females captured in late June carried 5.8 (5-7) embryos that had an average crown-rump length of 17.3 (7-27). One other individual had six placental scars. The average testicular length of 17 adult males obtained in June was 12.8 ( 10-15). Thirty-four of 50 females captured in July were pregnant; the average number of embiyos was 5.9 ( 2-9 ) , and their aver- age length was 12.1 (4-27). Four other females were lactating and another four had enlarged uteri, possibly denoting recent implantation. The average length of testes of 23 males obtained in July was 14.4 (10-18). A female captured north of Sundance on 15 August 1951 carried six fetuses and another obtained
southwest of Rapid City on 25 August 1952 contained eight (21 in crown-rump length). A subadult female taken in Wind Cave National Park on 1 Septem- ber 1968 was gravid, carrying three embryos that were 27 in cro\\'n-rump length. Many of the other above-men- tioned pregnant females were nearly ma- ture subadults. Juveniles are common in collections made from June through August, and undoubtedly in other peri- ods as well.
The process of pelage replacement in M. pennsylvaniciis is similar to that described for other microtines in the Black Hills. Adults molt from pale, faded winter pelage to fresh, dark sum- mer pelage in late June and July. All stages of molt, from commencement to completion, can be observed in adults taken on the same day at the same lo- cality in mid-summer. Juveniles were in a fine, dark fur that was replaced by subadult pelage. Developmental pelage replacement depends on chronology and rate of maturation of the individual; thus, concurrent existence of several different- aged litters tends to complicate interpre- tation of precise seasonal aspects of molt.
In the Black Hills, M. pennsylvaniciis are parasitized externally by ticks, Der- macentor andersoni Stiles, Ixodes spini- palpis Hadwen and Nuttall, and mem- bers of the Ixodes ochotonae-angustae complex, by laelaptid mites, Loehps mi- croti (Ewing) and Androlaelaps fahren- holzi (Berlese), and by a chigger, En- schoengastia setosa (Ewing).
J. A. Allen (1877:176) first reported meadow \'oles from the Black Hills area under the name Arvicola riparius, and later (1894b: 347) assigned Hills speci- mens to a new subspecies, insperatus. In his revision of the genus Microtus, V. Bailey (1900:20) listed Arvicola in- speratus in the synonomy of M. p. mo- destus, but later (1920:72) included Black Hills specimens under the name M. p. icahema described from eastern Montana and western North Dakota: R. M. Anderson (1943:92) returned in- speratus to subspecific rank, concluding
TURNER: MAMMALS Ol^^ THE BLACK HILLS
117
that icahenia was not distinct. S. Ander- son (1956) discussed subspeciation in the meadow vole in the western part of its range.
Examination of 31 adult male and 37 adult f(^male M. pennsylvanicu.s from the Black Hills revealed no secondary sexual variation in coloration. Males are significantly larger than females in lamb- doidal breadth (12.3±0.46, 11.9±0.37) and length of incisixe foramen (5.5± 0.31, 5.3±0.20), and are shghtly larger in most other dimensions. A pattern of variation in external and cranial dimen- sions is not discernible within the Hills but four specimens from west of Nemo (15.0 ±1.78) and six from northeast of Custer (15.1 ±1.28) are darker in tone of color on the average than 14 individ- uals from east of Buckhorn (18.6±:1.42), five from northeast of Sundance (18.3±: 2.17), 27 from south of Tinton (17.3± 1.87) and six from Beaver Creek Valley (16.5±1.18).
Microtus pennsylvanicus insperatus is evid(>ntly quite variable throughout its geographic range in external and cranial dimensions, but each character seems to \ary independently and no trends are ap- parent (Table 27). Meadow voles from the Black Hills are much darker in com- parison to specimens from Wyoming and Montana, but voles from Harding County, South Dakota, approach the color of those of the Hills in tone. Meas- urements additional to those in Table 27 for 68 adult M. pennsijlvanicus from the Black Hills are: percent red reflect- ance, 51. 2 ±0.24; percent green reflect- ance, 25.9 ±0.20; percent blue reflect- ance, 22.6 ±0.24; hind foot length, 20.8 ±3.83; weight, 48.6±7.44; interorbital constriction, 3.6 ±0.15; length of incisive foramen, 5.4±0.27; and length of maxil- lary tooth-row, 6.7±0.21.
Specimens examined ( 301 ) . — SOUTH DA- KOTA: Lawrence County: Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 4;
Table 27. — Geographic variation in selected color, and external and cranial measurements of adult
Microtus pennsylvanicus from the Northern Great Plains.
|
Number and sex of specimens averaged |
Tone of color |
Total length |
Tail length |
Ear length Greatest length of skull Zygomatic breadth |
Lambdoidal breadth |
Rostral length |
SkuU depth |
|
68(315,37$) Mean SD |
17.2 ±2.00 |
164.8 ±6.30 |
Black Hills, South Dakota and Wyoming 41.7 13.5 27.6 15.1 12.0 ±3.61 ±1.42 ±0.79 ±0.52 ±0.45 |
8.4 ±0.38 |
10.7 ±0.32 |
||
|
5(25,3$) Mean SD |
17.5 ±2.00 |
164.4 ±9.71 |
43.0 ±4.06 |
Harding County, South Dakota 12.8 27.4 15.6 ±1.30 ±1.11 ±0.41 |
12.3 ±0.27 |
8.3 ±0.54 |
10.6 ±0.24 |
|
7(45,3$) Mean SD |
19.1 ±2.65 |
172.1 ±4.02 |
44.3 ±2.75 |
Campbell County, Wyoming 13.4 27.5 14.8 ±2.30 ±0.81 ±0.50 |
11.5 ±0.70 |
8.6 ±0.41 |
10.8 ±0.22 |
|
29(16 5,13$ ) Mean SD |
19.3 ±2.32 |
168.3 ±7.97 |
49.1 ±2.98 |
Johnson County, Wyoming 13.0 27.4 ' 15.5 ±0.74 ±0.57 ±0.66 |
12.2 ±0.39 |
8.3 ±0.30 |
10.8 ±0.39 |
|
16(75,9$) Mean SD |
19.9 ±1.41 |
169.3 ±8.27 |
44.3 ±4.22 |
Sheridan County, Wyoming 13.9 27.9 15.4 ±0.81 ±0.62 ±0.54 |
12.4 ±0.34 |
8.3 ±0.26 |
11.0 ±0.40 |
|
10(45,6$) Mean SD |
19.8 ±2.08 |
162.5 ±4.35 |
49.3 ±2.11 |
Philips County, Montana 12.3 26.9 14.7 ±0.67 ±0.56 ±0.36 |
11.8 ±0.28 |
8.0 ±0.31 |
10.6 ±0.26 |
118
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
2 mi S Tinton, 6100 ft, 52; Roubiax Lake, 2 (NRW); 2 mi W Nemo, 4700 ft, 6. Penning- ton Countt/: Rapid City, 1 (NRW); Rapid Creek, 1.5 mi W Rochford, 3 (UMMZ); 2 mi W Oreville, 5500 ft, 1 (UMMZ); Beaver Creek, 4 mi N, 10.5 mi W Deerfield, 6400 ft. 18; Beaver Creek, 3 mi N, 9 mi W Deerfield, 6400 ft, 6; Castle Creek, 6500 ft, 7 (UMMZ); Bald Hills, Pactola, 3 (USNM); 6 mi S, 9.5 mi W Rapid City, 3; Sheridan Lake, 2 (NRW); Spring Creek, south Rapid City, 1 (MHM); 4 mi NW Hill City, 2 (UMMZ); 20 mi N Elk Mountain, 6000 ft, 5 (USNM); Moon, 22 mi W Hill City, 6200 ft, 7; Ditch Creek, 14 mi W Hill City, 6400 ft, 1; Hill City, 1 (AMNH); near Palmer Culch, 1 ( UMMZ ) ; Palmer Gulch,
3 mi SE Hill City, 5300-5400 ft, 25 (UMMZ); Nelson's Place, 4 mi SE Hill Citv, 5300-5400 ft, 10 (UMMZ); 16 mi NW Custer, 3 (UMMZ); Horse Thief Lake, near Mount Rushmore, 1 (NRW); imspecified locality, 3 (UMMZ). Custer County: Palmer Gulch, 8 mi SE Hill City, 5300 ft, 14 (FMNH); 3.5 mi S, 0.5 mi W Kevstone, 5000 ft, 12; 0.5 mi E Sylvan Lake, 6250 ft, 1 (UMMZ); 5.75 mi N 5.75 mi E Custer, 5220 ft, 13; Custer, 5 (AMNH); Harney National Forest, 4 mi E Custer, 2 (UMMZ); 3 mi N Pringle, 5000 ft, 3 (UMMZ); Lightning Creek, 8 mi SW Custer, 5100 ft, 1 (UMMZ); Beaver Creek Canyon, Wind Cave National Park, 4200 ft, 8; Head- quarters, Wind Cave National Park, 4100 ft, 1; Wind Cave Canyon, Wind Cave National Park, 4100 ft, 1; Bull Springs, 2 mi N, 9 mi W Custer, 9 (UMMZ).
WYOMING: Crook County: Bear Lodge Mountains, 2 (USNM); Bear Lodge Mountains, 6.5 mi SSE Alva, 2 (UMMZ); 15 mi N Sun- dance, 5500 ft, 3; 15 mi ENE Sundance, 3825 ft, 6; 3 mi NW Sundance, 5900 ft, 1; 1.5 mi NW Sundance, 5000 ft, 4; Sundance, 15 (USNM); Ratdesnake Creek, 6000 ft, 1 (USNM); Sand Creek, 3750 ft, 2 (USNM). Weston County: 1.5 mi E Buckhorn, 6150 ft, 25; Newcastle, 2 (USNM).
Additional recorrf.s.— SOUTH DAKOTA: Lawrence County: Spearfish (BB). Penning- ton County: (BB). Custer County: French Creek, 3 mi E Custer (Stebler, 1939:390).
Ondatra zibethicus cinnamominus
(Hollister)
MUSKRAT
Fiber zibethicus cinnamominus Hollister, 1910, Proc. Biol. Soc. Washington, 23:125, 2 Sep- tember (type locality, Wakeeney, Trego Co., Kansas).
Ondatra zibethicus cinnamomina — Miller, 1912, Bull. U. S. Nat. Mus., 79:232, 31 Decem- ber.
The muskrat is cliaracteristic of the prairie sloughs, marshes, and drainage
systems of the Northern Great Plains and is common on nearly all streams that originate in the Black Hills. Although few specimens of this semiaquatic micro- tine are available, I have observed musk- rats throughout the Hills. Dens and sub- merged trailways interlace the shorelines of lakes and reservoirs, and penetrate banks of all water courses of the region. In the past, Ondatra undoubtedly has been one of the most commercially im- portant fur-bearers sought by trappers in the Black Hills, but more recently, muskrat pelts ha\'e fetched relatively poor prices. Burrows excavated in dams and irrigation ditches cause extensive damage to these structures and lend a negatixe dimension to the economic im- portance of this species.
Compared to O. z. osoijoosensis to the west in Wyoming, O. z. cinnamominus is smaller cranially, with a notably shorter maxillary tooth-row, and is paler in color, being more reddish or buflPy instead of brownish (Long, 1965:662). The species was first reported from the Black Hills by Bailey (1888:445).
Reproducti\'e and natural history data are lacking for this microtine in western South Dakota, but in northern Nebraska, the muskrat breeds from late April to early September — adult females produce one to four litters per year, averaging about six voung per litter (Jones, 1964: 235).
Specimens examined (24). — SOUTH DA- KOTA: Lawrence County: Little Spearfish Canyon, Savoy, 4 (USNM). Pennington County: Rapid City, 2 (1 USNM); Diamond S Ranch, near Rapid City, 2 (UMMZ); Spring Creek, south of Rapid City, 2 (MHM); Tiger- ville, near Hill Citv, 1 (USNM); Hill Cit\-, 3 (AMNH); Harney National Forest, 1 (UMMZ). Custer County: Custer, 8 (7 AMNH, 1 FMNH). Fall River County: 1 mi N Hot Springs, 1 (UW).
Additional records.— SOVTH DAKOTA: Custer County: French Creek, 3 mi E Custer (Stebler, 1939:390). Fall River County: Hot Springs (Woodward, 1930:109).
Family MURIDAE— Old World Rats and Mice
The two species of minids occurring in the Black Hills are commensals of man,
TURNER: MAMMALS OF THE BLACK HILLS
119
and were introduced into North America from the Old World in the mid-eigh- teenth century.
Rattus norvegicus (Berkenhout) Norway Rat
Mas norvegicus Berkenhout, 1769, Outlines of the natural history of Great Britain and Ireland, 1:5 ( t\'pe locality, England, where the species was introduced from Asia, prob- ably \'ia Continental Europe, in the early ITOO's).
Rattus norvegicus — Hollister, 1916, Proc. Biol. Soc. Washington, 29:126, 6 June.
The Norway rat was introduced to tlie colonies of eastern United States from the Old World about 1775, escaping from ships at various seaports (Silver, 1941: 2). I do not know the history of its im- migration into the Black Hills region, but the species presumably was intro- duced at rixer towns and forts along the Missouri Valley in the 1850's and sub- sequently dispersed in a manner similar to Mus musculus, as described in the following account.
I know of only two actual specimens from the study area, but Over and Churchill (1945:45) indicated that these rats were found occasionally in the Black Hills, in and around habitations of man.
Specimens examined (2). — SOUTH DA- KOTA: Pennington County: Rapid City, 2 (SDMT).
Additional /T'corc/.— SOUTH DAKOTA: "Black Hills" (Over and Churchill, 1945:45).
Mus musculus Linnaeus House Mouse
[Mus] musculus Linnaeus, 1758, Systema nat- urae, ed. 10, 1:62 (type locality restricted to Upsala, Sweden, by Thomas, Proc. Zool. Soc. London, p. 147, March 1911).
The house mouse is a common, often abundant, rodent that frequently is in- jurious to man and to the native fauna. It resides in and around human habita- tions at lower elevations in the Black Hills. For example, on 13 June 1967, sev- eral dozen individuals were killed with .22 dust-shot in grain bins near Minne- kahta. Fall Ri\er' County. Wild popula- tions occasionallv are encountered.
This rodent was introduced from Europe in early times and has since spread over most of temperate and tropical North America. I am unfamiliar with the history of its dispersal to the Black Hills, but Mus niuscuhis presum- ably was introduced in the plains area throughout the Mississippi and Missouri river valleys from river boats to wharves of forts and trading posts; thence, it probably spread into the study area in company with early military expeditions and the establishment of settlements. Hayden (1859:710) mentioned that both the house mouse and Norway rat were abundant at all of the fur trading posts on the Missouri River in the 1850's.
Because two subspecific stocks were introduced into North America ( Schwarz and Schwarz, 1943:65), I choose not to assign a subspecific identity to speci- mens from the Black Hills.
A female taken southeast of Spearfish in mid-August evidenced six placental scars, whereas another obtained in Wind Cave National Park in late August showed no signs of reproductive activity. Testicular lengths of two males obtained in June from near Minnekahta were 5.
Specimens examined (25). — SOUTH DA- KOTA: Lawrence County: 1.5 mi S, 0.5 mi E Spearfish, 3800 ft, 1; 8 mi SW Lead, 1 (UW). Meade County: Black Hawk, 1 (AMNH). Pennington County: 4 mi N Rapid Cit>', 3 (AMNH); Rapid City, 12 (10 NRW, 2 SDMT); Glendale, 1 (AMNH). Custer County: Squaw [Grace Coolidge] Creek, 1 (AMNH); 2.5 mi N, 5 mi E Wind Cave, Wind Cave National Park, 3700 ft, 1; Wind Cave Can- yon, Wind Cave National Park, 4100 ft, 1; Wind Cave National Park, 1 (USNM). Fall River County: 0.5 mi S, 1.5 mi W Minnekahta, 4200 ft, 2.
Additional record.—SOVTH DAKOTA: Lawrence County: Spearfish (BB).
Family ZAPODIDAE— Jumping Mice
Zapodids, represented in the Black Hills region by one species, are salta- torial rodents that abide in areas of lush vegetation such as wet meadows or streamside communities. Aridity is the major factor restricting the distribution of jumping mice on the Northern Great Plains.
120
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Zapus hudsonius campestris Preble
Meadow Jumping Mouse
Zapus hudsonius campestris Preble, 1899, N. Amer. Fauna, 15:20, 8 August (type lo- cality, Bear Lodge Mountains, Crook Co., Wyoming).
Although specimens are absent from Meade and Fall River counties, the mea- dow jumping mouse is common through- out the study area. It most frequently occurs in riparian communities along small streams in valley meadows, or in open, moist habitats within coniferous forests that support a low undergrowth of forbs and grasses. Zapus hudsonius occurs in suitable habitats on the ad- jacent plains, but is most frequent in the Black Hills up to 6500 feet. This species hibernates in winter, and all specimens at hand were taken in warmer months; earliest and latest seasonal rec- ords are 4 June and 27 August, respec- tively.
Jumping mice collected in this study were trapped along drainage systems, in dense grasses and sedges that were in- terspersed with deciduous shrubs and fallen logs of aspen, birch, willow, white spruce, and ponderosa pine. Most of the specimens from the Wyoming part of the Hills, and those individuals from Little Spearfish, Big Spearfish, and Beaver Creek canyons were taken under these general conditions. Similarly, five fe- males and three males were trapped in a large valley of approximately one mile in length located southwest of Keystone. A stream, with many small tributaries, flowed through the center of the valley and created a semimarsh situation. Lush vegetation was mainly of tall grasses and some very dense patches of clover; de- ciduous shrubs bordered the stream.
Forty-two individuals (22 males and 20 females) were taken in the lowland habitat of Grizzly and Iron creeks, north- east of Custer, in mid-June 1967. Ri- parian habitat was composed of grasses, spearmint, yellow wood sorrel (Oxalis stricta), stinging nettle, and ferns; drift- wood, woodpiles, and shrubs were dis- persed throughout. Eleven male and
three female jumping mice were trapped west of Hill City, in a marshy lowland bordering Ditch Creek. Small pools of water were interlaced with hummocks of soil that supported growths of liverworts and mosses. Grasses, sedges, and white spruce also were predominant in the immediate area. Additional specimens were obtained under similar conditions near the east end of Deerfield Lake and along Goldrun Creek, southeast of Deer- field. Representatives of Z. /k campestris taken near Nemo, Lawrence County, were trapped along the banks of Box- elder Creek. In these various habitats, Sorex cinereus, Peromysctis manictilatus, Microtus lon0catidus, M. pennsyJvani- cus, and Eutamias minimus were trapped in association with Z. hudsonius.
Several jumping mice were collected in non-riparian habitats. A male from 14 mi W Hill City was trapped at the base of a large rock on a dry wooded slope that was far removed from a source of standing water. A male, from the same localitv, was shot in late afternoon as it ran from beneath the branches of a partialh' hollow log that was situated in a dry and rocky meadow. Another male from near Nemo was trapped on a dry hillside among ponderosa pine trees and granite boulders.
About 20:30 hrs (MDT) on 1 August 1967, I heard rustling in a stand of slough grass (Beckmannia syzigachne) growing along the shore of the Savoy Reservoir, Lawrence County. A jump- ing mouse was observed climbing an already-inclined stem of grass, thus rid- ing the stem to the ground. Straddling the bent stem and holding it close to the ground, the mouse systematically ro- tated the rachis and removed the seeds. Within about five minutes, the seed- laden head of the grass was bare, and the mouse departed carrying seeds in its cheeks, which were noticeably distended.
Fourteen of 31 females taken in the latter half of June carried 5.8 (4-7) em- bryos that were 13.1 (3-18) in crown- rump length; four others had 5.5 (4-7) placental scars. Average testicular length
TURNER: MAMMALS OF THE BLACK HILLS
121
of 38 males obtained in tliis period was 6.7 (4-10). In July, six of 2.3 females showed no signs of reproductive activity, but 11 pregnant females carried 6.1 (5-8) embr\os \\'hich were 12.9 (3-20) in crown-rump length. The uteri of six other indixiduals had 3.8 (2-5) placental scars. Average length of testes of 13 males obtained in July was 6.1 (4-8). Two of six females captured early in Au- gust had 5.5 (5-6) placental scars; none was pregnant. Testicular length of six males obtained at this time was 5.7 (4-7) . Most females that exhibited placental scars also were lactating.
Three young Zajnis were captured northeast of Custer in late June. Sub- adults, which are brighter in color than adults and generally lack a distinctive mid-dorsal stripe, are not uncommon throughout the warm months, until at least mid-August. A female li\e-trapped on 25 July 1967 at the last-mentioned site gave birth to seven still-born young during the ensuing night; external meas- urements of these newborn voung are: total length, 38.3 (37-39); tail length, 10.8 (9-13); and hind foot length, 5.0 (5).
The process of annual molt occurs in adults from mid- to late summer. For example, five males taken from 27 July to 2 August in Little Spearfish Canyon displayed bright-colored tips of new hairs projecting from beneath the old dull pelage of the dorsum. Two females from southwest of Rapid City and two males from north of Sundance (all cap- tured in mid-August ) were in completely new pelage.
Robertson (1971) recently analyzed variation in populations of Z. hudsonius on the Great Plains. Because his studies were concurrent with my own, and be- cause we had a mutual exchange of in- formation, I rely upon his results herein. Jumping mice exhibit clinal variation on the plains northwestward from eastern Kansas, becoming progressively darker in coloration and tending toward longer, narrower, deeper skulls. Robertson's findings (1971:20-21) agree with those
of Krutzsch (1954:441), indicating that Z. h. campestris, originally described from the Black Hills, contrasts with Z. h. pallidiis to the east in South Dakota in being conspicuously darker and axerag- ing larger in external and cranial meas- urements.
Secondary sexual dimorphism is not evident in jumping mice from the study area. However, patterns of variation in size within the Hills are discernible. In- dividuals from Pennington County are larger than those from other Black Hills populations in occipitonasal length, zygo- matic length, length of palatine foramen, and cranial depth, but are smaller in length of maxillary tooth-row, zygomatic breadth, mastoid breadth and width of palatine foramen. These dimensional re- lationships are reciprocal when compar- ing representatives from Lawrence County with those of other areas (Rob- ertson, 1971:16-17). Physiographic ir- regularity of the Hills may contribute to this morphological diversity. Coloration shows no readily recognizable trends in the Black Hills.
Selected external and cranial meas- urements of 20 adult males, followed by those of 20 adult females, are (Robert- son, 1971, Table 1): total length, 125.3+5.43, 126.2+6.86; hind foot length, 29.6 + 1.47, 29.5+1.64; ear length, 14.4+1.70, 13.4 + 1.60; occipitonasal length, 23.5 + 0.54, 23.2+0.46; zygomatic length, 9.5+0.25, 9.5 + 0.28; zygomatic breadth, 11.4+0.35, 11.3+0.39; mastoid breadth, 10.1+0.38, 10.0+0.45; inter- orbital constriction, 4.2 + 0.22, 4.3+0.20; cranial depth, 9.1+0.16, 9.2+0.27; palate length, 3.5 + 0.24, 3.5+0.23; length of maxillary tooth-row, 3.6 + 0.17, 3.5+ 0.15; length of palatine foramen, 4.6+ 0.29, 4.4 + 0.22; width of palatine fora- men, 2.2+0.14, 2.1ih0.14.
Specimens examined (207).— SOUTH DA- KOTA: Lawrence CounUj: Big Spearfish Can- yon, 9 mi S, 3 mi W Spearfish, 5000 ft, 9; Little Spearfish Canyon, 2 mi S, 10 mi W Lead, 5800 ft, 14; 2 mi S Tinton, 6100 ft, 7; 3 mi W Nemo, 4800 ft, 1; 2 mi W Nemo, 4700 ft, 9; Nemo, 4700 ft, 1. Pcunin^fon Conniij: 1 mi S, 8.5 mi W Rapid City, 2; Beaver Creek,
122
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
4 mi N, 10.5 mi W Deerfield, 6400 ft, 10; Beaver Creek, 3 mi N, 9 mi W Deerfield, 6400 ft, 1; Goldrun Creek, 1 mi S, 2 mi E Deer- field, 6400 ft, 3; Rapid Creek, 2 mi W Pactola, 4800 ft, 3 (UMMZ); Castle Rock, R. 2 E, T. 1 N, 6500 ft, 3 (UMMZ); Ditch Creek, 14 mi W Hill City, 6400 ft, 10; Ditch Creek, 13 mi W Hill City, 6400 ft, 4; Nelson's Place,
3 mi SE Hill Cit\% 6 ( UMMZ ) ; Palmer Gulch,
4 mi SE Hill City, 6 (UMMZ); mispecified locality, 1 (UMMZ). Citster County: 3.5 mi S, 0.5 mi W Kevstone, 5000 ft, 8; Palmer Gulch, 8 mi SE Hill City, 9 (FMNH); 5.75 mi N, 5.75 mi E Custer, 5220 ft, 42; Custer, 5 (2 AMNH, 3 USNM); Wind Cave National Park, 1 (WCNP); Game Ranch, Cold Spring Creek, Wind Cave National Park, 2 (UMMZ); Bea\'er Creek Canvon, Wind Cave National Park, 4200 ft, 9 (I'UMMZ).
WYOMING: Crook County: Bear Lodge Mountains, 6 ( USNM ) ; Bear Lodge Mountains, 6.5 mi SSE Alva, 1 (UMMZ); Bear Lodge Mountains, Warren Peak Lookout, 6000 ft, 2 (USNM); Dexils Tower, flood plain Belle Fourche River, 3350 ft, 1 (USNM); 15 mi N Sundance, 2; 3 mi NW Sundance, 5900 ft, 19; Sundance, 3 (USNM). Weston County: 1.5 mi E Buckhorn, 6150 ft, 7.
Additional recor J.s.— SOUTH DAKOTA: Latcience County: Spearfish (BB). Penning- ton County: Rochford (BB). Custer County: Bull Springs (Bole and Moulthrop, 1942:167).
Family ERETHIZONIDAE— Porcupines
Only one genus, with one species, of this large, quilled rodent occurs in the Black Hills region.
Erethizon dorsatum bruneri Swenk Porcupine
Erethizon epixanthum hruneri Swenk, 1916, Univ. Nebraska Studies, 16:117, 21 Novem- ber (type locality, 3 mi E Mitchell, Scott's Bluff Co., Nebraska).
Erethizon dorsatum hrw^eri — Anderson, 1947, Bull. Nat. Mus. Canada, 102:173, 24 Janu- ary.
Erethizon dorsatum occurs on the Great Plains and adjacent montane areas at sites where trees are available locally. In the Black Hills, porcupines have a wide altitudinal distribution, occurring from the boreal cap, down through the transition zone, and out onto the sur- rounding prairie. This rodent is active the year around. Although common in the Hills, specimens of Erethizon are
lacking from Fall River and Weston counties.
Erethizon dorsatum hruneri differs from E. d. epixanthum, which occurs to the west in Wyoming, in ha\'ing smaller external and cranial dimensions, arched frontals, and auditory bullae that are less inflated (Long, 1965:669).
Porcupines usually were encountered along the shoulders of highways in eve- ning or early morning, and many speci- mens in collections were taken as road- kills. Due to heavy damage inflicted by this species upon potentially harvestable timber, particularly young pine trees, the U. S. Forest Service maintains a porcu- pine control program in the Hills. Trees with tops and branches barked indicate the abundance and widespread distribu- tion of Erethizon in the study area.
Sixteen indi^'iduals were collected along Little Spearfish Canyon in late July and early August of 1967, generally along roadways at dusk. On t\\'o occa- sions, juvenile males were taken in com- pany with large adult males. One procu- pine was shot as it persistently chewed a hole in a camper's tent; bacon grease had been spilled on the tent and even repeated attacks on the porcupine with a club could not drive the rodent awa\'. Another indi\idual was shot at mid-day as it roosted in the top of a pine at the bottom of Big Spearfish Canyon.
Rocky bluffs are used as den sites by Erethizon; dens are evident from ac- cumulations of feces and discarded quills. Porcupine scats are in the form of pellets, which are often interconnected by strands of undigested vegetation. Fre- quently dens of Erethizon are associated with those of Marmota faviventris.
A female taken near Rockerville on 29 March 1946 contained one embryo. Females obtained in July and August each showed a single placental scar in the uterine horns. A lactating female was collected northeast of Custer on 17 June 1967, and juveniles were common from July through September. The average length of testes of four males taken in August was 44.2 (40-57). Ju\enile pelage
TURNER: MAMMALS OF THE BLACK HILLS
123
consists of fine soft blackish underfur and long yellowish guard hairs, as com- pared to the rather coarse underfur and whitish guard hairs of adults. The quills are less resilient than tliose of adults, and also possess a yellowish tinge.
Specimens examined (52). — SOUTH DA- KOTA: Lawrence County: Big Spearfish Can- yon, 3 mi W Spearfish, 4200 ft, 1; Tinton, 5900 ft, 1; 2 mi S Tinton, 6100 ft, 1; Little Spear- fish Canyon, Roughlock Falls, 9 mi W Lead, 5400 ft, 1; Little Spearfish Canyon, 1 mi S, 9 mi W Lead, 5400 ft, 5; Little Spearfish Can- yon, 2 mi S, 10 mi W Lead, 5800 ft, 7; Little Spearfish Canyon, 6.2 mi SW Lead, 6000 ft, 1; Little Spearfish Canyon, 12 mi S, 14 mi W Lead, 6200 ft, 2; Bogus Jim Creek, 1 (SDMT). Meade County: Vanocker Canyon, 3 mi S, 0.5 mi W Sturgis, 4400 ft, 1. Pennington County: 1.5 mi N Silver City, 1 (SDMT); 0.4 mi SW Silver City, 1 (SDMT); Rapid City, 1 (NRW); Bea\'er Creek, 3 mi N, 9 mi W Deerfield, 6500 ft, 1; 1 mi NW Deerfield, 1; Deerfield, 1 (SDMT); Spring Creek Canyon, 4 mi ENE Rockerville, 3600 ft, 2 (UMMZ); Ditch Creek, 14 mi W Hill City, 6400 ft, 1; 19 mi W, 3 mi S Hill City, 1; Palmer Gulch, 4 mi SE Hill City, 5300 ft, 1 (UMMZ); un- specified locality, 2 (SDSU). Custer County: RoJjy Canyon, 2 mi N, 22 mi W Custer, 5200 ft, 1; 5.75 mi N, 5.75 mi E Custer, 5220 ft, 1; Shirttail Canyon, 2 mi S Wind Cave, Wind Cave National Park, 1 (WCNP); Dewey, 11 (USNM).
WYOMING: Crook County: 15 mi N Sun- dance, 1; 15 mi ENE Sundance, 1; Sundance, Bear Lodge Mountains, 6000 ft, 1 (USNM). Weston County: 1.5 mi E Buckhorn, 6150 ft, 1.
Additional records.— SOUTH DAKOTA: Custer County: "vicinity of Otis" (Stebler, 19.39:389). WYOMING: Weston County: "east of Newcastle" (Stebler, 1939:389).
ORDER CARiNIVORA— Carnivores
A rather diverse group of mammals from the diminutive ermine to the robust grizzly bear, comprise the carnivorous fauna (nine genera and 15 species) in the Black Hills within historic time. The statuses of an additional seven species (si.x genera) in the Hills region remain uncertain.
Representatives of all five families of Carnivora that occur in North America either are indigenous to the Black Hills or recently have been extirpated by man. Large carni\'ores tend to interfere with man's pastoral enterprises, and as a re-
sult, the gray wolf, black bear, grizzly bear, mountain lion and lynx either no longer occur in the Hills, or are rare. Coyotes, badgers, and bobcats also must endure the attempts of humans to deci- mate their populations.
Family CANIDAE— Coyotes, Wolves, and Foxes
This family of doglike mammals is represented in the area under study by two genera and three species. Two addi- tional canids, the swift fox and gray fox, occur in the general vicinity and may oc- casionally intrude into the Black Hills.
Canis latrans latrans Say
Coyote
Cams latrans Say, 1823, in Long, Accoimt of an expedition from Pittsburgh to the Rocky Mountains . . ., 1:168 (footnote), (type locality. Engineer Cantonment [approxi- mately 2 mi E Ft. Calhoun], Washington Co., Nebraska).
The coyote is known from much of North America, and is well distributed throughout the Black Hills. At least 200 individuals were poisoned on the Sheid- lev Cattle Companv range near Rapid city in 1894 (Bailey, 1907:11), 1165 coyotes were killed in the Bear Lodge Mountains in 1907 (Bailey, 1908:6), and reports of District Rangers, Black Hills National Forest (U. S. Biological Survey files), indicate that approximately 2680 coyotes were killed in that area in the period from 1926 to 1935. From 1962 to 1968, bounties on 19.30 coyotes were paid by the counties that comprise the South Dakotan portion of the Hills ( rec- ords from South Dakota Dept. Game, Fish and Parks), and extensive use of 1080 poison (sodium monoHuoracetate) has further reduced the population in the eastern and southern sections of the Black Hills (A. H. Richardson, pers. com.). In spite of these attempted con- trol measures, Canis latrans seems to be increasing in abundance in the northern Hills, especially along the South Dakota- W\'oming border (A. W. Jones, pers. com. ) .
124
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Compared to Canis latrans Jestes, which occurs to the west in Wyoming, C. /. latrans is slightly smaller externally and cranially, and the upper parts are much paler in color (Long, 1965:674).
Evidently, coyotes increasingly in- vaded the Black Hills as gray wolves were extirpated by man, and became much more abundant as the latter species was reduced in numbers. Grinnell ( 1875: 79-80) stated: "The coyote was found in considerable numbers on the plains, and was especially abundant among the elevated table-lands that were crossed just before reaching the Black Hills. After penetrating into the hills proper, however, I did not see a single specimen until I left them for the Big Cheyenne [Cheyenne River], when I again noticed coyotes in numbers. In the Black Hills, this species would seem to be replaced by the preceding [gray wolf]." In 1894, W. W. Granger indirectlv referred to a change in relative abundance of the two species when he noted that gray wolves were "not uncommon," and coy- otes were "common" in the Black Hills (J. A. Allen, 1895a: 274). Coyotes con- tinued to increase in numbers in the Hills after the turn of the century, whereas the gray wolf was eliminated shortly thereafter. Conversely, the de- cline in ungulate populations, important as a source of food for coyotes, and in- creased conflict with the agricultural pur- suits of man, caused a reduction in the number of C. latrans on the plains.
I often have observed C latram near dusk and shortly after dawn in the various prairie dog towns in Wind Cave National Park. On 25 July 1968, I watched a large adult "mousing" along Highland Creek for several hours in mid- morning. A litter was raised in Beaver Creek Canyon that summer and the pups were vocally active each evening. On 27 August 1968, an adult coyote and three young were observed pursuing a doe pronghorn in Shirttail Canyon. The fate of the pronghorn remains unknown as all animals (quickly passed from view; however, no evidence of its capture could
be found in the subsequent inspection along the route of the chase.
Reproductive data applicable to the Black Hills coyote population were ob- tained from the U. S. Biological Survey files. Two females taken near Custer in March 1944 carried four and sexen embryos, respectively; nine females ob- tained in the Hills in late April and early May had an average of 6.4 (4-7) embryos each. Two adult females (one with se\'en suckling pups and the other with four embryos) were captured in Wind Cave National Park late in April 1918 by Troy C. Beach. Thus, coyotes in the Black Hills, as elsewhere on the northern plains, apparently breed in Feb- ruary and March, bearing litters in April and May (Mengel, 1971:325).
Stomachs of five coyotes, trapped in Wind Cave National Park in early June 1924, contained the remains of domestic chicken, domestic lamb, and cottontail rabbit (Sylvilagus sp.).
Specimens examined (19). — SOUTH DA- KOTA: Pennington County: 4 mi S, 14 mi W Hill City, 2. Custer Countij: 18 mi NE Dewev, 4 (USNM); 12 mi NE Dewev, 1 (USNM); 4 mi N Dewey, 1 (USNM); 5 mi NW Wind Cave, 1 (WCNP); Wind Cave Canyon, Wind Cave National Park, 1 (WCNP); Shirttail Can- yon, Wind Cave National Park, 2 (WCNP); 3 mi SE Wind Cave, 1 (WCNP). Fall River Count!/: 5 mi W Minnekahta, 1.
WYOMING: Crook County: Bear Lodge Mountains, 2 (USNM); Grand Canyon, Sun- dance, 1 (USNM); Rattlesnake Canyon, Sun- dance, 1 (USNM).
UNSPECIFIED LOCALITY: Black Hills, 1 (SDSU).
Additional records (USES card files unless otherwise noted).— SOUTH DAKOTA: Law- rence County: Spearfish; Lead. Meade County: Piedmont. Pennington County: Rochford; Deerfield; 17 mi NW Custer. Custer County: Bull Springs, 14 mi NW Custer (Lamster, 1943: 1); Custer; 9 mi W Pringle; Pringle; Dewey; 18 mi W Custer; unspecified locality, 1 (Jack- son, 1951:262). Fall River County: ' Hot Springs; Minnekahta.
Canis lupus irremotus Goldman
Gray Wolf
Canis lupus irremotus Goldman, 1937, Jour. Nhmun., 18:41, 14 February (type locality, Red Lodge, Carbon Co., Montana).
TURNER: MAMMALS OF THE BLACK HILLS
125
The gray \vo\i formerly occurred throughout much of the United States. Historical reports indicate that Conis lupus pre\iously was abundant and widely distributed on the Great Plains and in the Black Hills. Extirpation of the vast herds of bison destroyed the main food source of the \\'olf and this, along with trapping and poisoning of wolves for protection of livestock, for fur, and for bounties, finally led to the elimina- tion of this species from major areas of its former range.
Conis lupus irremottis differs from C /. nuhilus to the south and east in being larger, paler, and narrower in the frontal region, especially relative to the width of the rostrum (Long, 1965:677). Goldman (1944:448-449) noted that specimens from the Black Hills are vari- able and grade toward nubilus.
Grinnell (1875:79) wrote of this spe- cies as follows: "I found the gray wolf one of the most common animals in the Black Hills, and hardly a day passed without my seeing several individuals of this species. They were generally ob- served singly or by twos and threes, sneaking along the mountain sides or crossing the narrow valleys." He indi- cated that in the Black Hills, C. latrans (see account of this species) was re- placed by C. lupns. Dodge (1876:133) noted : "The wolves are of enormous size, a very dark gray in color, and in consid- erable numbers." From the spring of 1895 to the autumn of 1897, about 500 gray wolves were killed on the range of the Ames Cattle Company (54.35 square miles) in northeastern Crook Count}', Wyoming (Long, 1965:676). J. T. Craig, manager of Western Ranches Limited, Belle Foiu"che, South Dakota, filed a re- port with the U. S. Biological Survey in 1896, indicating that C. lupus was re- sponsible for a three percent loss of calves, one percent loss of adult cattle, and five percent loss of colts annually (Young, 1946a: 122). G. E. Lemmon, manager of the Sheidley Cattle Com- pany, Rapid City, South Dakota noted that 80 to 100 gray wolves were poisoned
on ihc range of this company in 1896 (Bailey, 1907:11). Near Newcastle, wolves were still numerous enough to be destructive to li\estock in 1905 and 1906 (Bailey, 1907:10). Evidently, the gray wolf was already becoming more scarce at this time, whereas the coyote was becoming more abundant in the Black Hills (see Allen, 1895a: 274).
Decimation of C. lupus in the Hills continued during the early part of the twentieth century (U. S. Biological Sur- vey files, unless otherwise noted ) . About 925 gray wolves were obtained by the U. S. Forest Service in the Bear Lodge Mountains and surrounding portions of Crook County, Wyoming, in 1907 (Bailey, 1908:6). Bounties were paid in 1911 on 55 wolves by counties that com- prise the South Dakotan section of the Hills (records from South Dakota Dept. Game, Fish, and Parks). Eight wolves were destroyed in 1915, and five more were killed in 1916. A pregnant female that carried four embryos was taken 18 mi NE Dewey on 25 March 1917. A male was killed 15 mi SE Dewey on 10 November 1920. Three indi\'iduals were shot on 23 December 1926, and two were taken on 11 January 1928.
One of the famous American rene- gades was the "Custer Wolf" which ranged in the vicinity of Custer for a period of seven years (Young, 1946a: 68, photo opposite). This large male had a known range of 40 by 60 miles (Seton, 1929a: 257) and was credited with killing $25,000 worth of livestock during this period. It avoided capture despite a bounty of $500 off^ered for its scalp. A Federal Hunter, H. P. Williams, concen- trated entirely upon the "Custer Wolf" from March to October 1920 before fi- nally succeeding in killing it (Young, 1944:277). The last specimen from the Black Hills known to me is one reported by the Distiict Forester in Harney Na- tional Forest, 30 January 1934 (U. S. Biological Survey files ) . On 4 July 1967, I found a mandible of C. lupus in a woodrat nest at the rear of Davenport Cave, southwest of Sturgis. Currently,
126
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
the former range of C. lupus in the Black Hills has been pre-empted by Canis latrans.
Specimens examined (3). — SOUTH DA- KOTA: Meade County: Da\'enport Cave, 3 mi S, 1.5 mi W Sturgis, 4400 ft, 1. Custer County: 12 mi NE Dewey, 1 (USNM).
WYOMING: Crook County: Sand Creek Canyon, 1 (USNM).
Additional records.— SOUTH DAKOTA: Custer County: 18 mi NE Dewey (USES files); Dewey (Goldman, 1944:445). Fall River County: 15 mi SE Dewey (USES card files). WYOMING: Crook County: northeast- ern portion of county (Long, 1965:676).
Vulpes vulpes regalis Merriam Red Fox
Vulpes regalis Merriam, 1900, Proc. Washing- ton Acad. Sci., 2:672, 28 December (type locality. Elk River, Sherburne Co., Minne- sota ) .
Vulpes fulva regalis — E. Bailey, 1929, Jour. Mamm., 10:157, 9 May.
The red fox is widely distributed and relatively common in the eastern Great Plains, and occurs sparingly in suitable habitat in the eastern and southern por- tions of the Black Hills. Grinnell (1875: 96) observed three red foxes, an old female and two young that inhabited a burrow on the edge of a prairie dog town along the northern fringe of the Black Hills in 1874. Bailey (1888:432) witnessed "a few tracks" of this species in the snow in October 1887, and on 20 July of the following year obtained a female near Custer, noting a field report to the U. S. Biological Survey that this species was "common" in that vicinity. A male was taken by Merritt Gary near Elk Mountain on 16 October 1903.
Game Biologists A. H. Richardson and L. Petersen, and Game Warden E. L. Woods, indicated to me that Vulpes vulpes was relatively unknown in the Black Hills five to 10 years ago, but that since 1965 it has increased noticeably in the region (pers. com.). These natural- ists felt that the coyote population has decreased considerably in some areas of the Black Hills because of extensive use of 1080 poison, and that the red fox is replacing the coyote in the southern and
eastern sections (see additional records for specimens that recently have come to the attention of these men).
A red fox recently was taken along the southern periphery of the Black Hills at a locality 10 mi N Rumford, Fall Ri\'er Gounty (Jones and Henderson, 1963: 283). 'From 1962 to 1968, bounties on 1079 V. vulpes have been paid by those counties that comprise the South Da- kotan portion of the Black Hills ( records from South Dakota Dept. Game, Fish, and Parks ) .
Vulpes vulpes regalis diflFers from V. V. macroura of western Wyoming in being larger and more golden-reddish instead of yellowish in color (Long, 1965:679). As other recent authors, I follow Ghurcher ( 1959 ) in using the spe- cific name Vulpes vulpes (Linnaeus) for American red foxes.
Specimens examined (2). — SOUTH DA- KOTA: Custer County: Custer, 1 (USNM); Elk Mountain, 1 (USNM).
Additional records (A. H. Richardson, L. Petersen, and E. L. Woods, pers. com.). — SOUTH DAKOTA: Lawrence County: near Nemo, 1. Pennington County: 7 mi NW Rapid City, 1. Custer County: 2 mi S Custer, 1; 2 mi W Hermosa, 1. Fall River County: 2 mi N Hot Springs, 1; 2 mi S Hot Springs, 1.
Family URSIDAE— Bears
The large omnivorous ursids are rep- resented in the Black Hills by one genus and two species.
Ursus americanus americanus Pallas Black Bear
Ursus americanus Pallas, 1780, . . . Spicilegia zoologica . . ., fasc. 14.5 (type locality, eastern North x\merica).
Although I have examined but a single specimen of this species, historical accounts as well as recent newspaper accounts and reports by National Forest Service personnel, indicate the presence of this bear in the Black Hills.
Dodge (1876:132) wrote of bears in the Black Hills as follows: "The country in this part of the Hills [along Rapid and Boxelder Greeks in Pennington Gounty] is full of bear sign. In some places almost
TURNER: MAMMALS OF THE BLACK HILLS
127
a quarter of an acre will be rooted up as if by hogs; small thickets of berry-bear- ing bushes are torn and broken; ant-hills are dug into and huge logs turned over by this omnivorous monster in search of his food." He (1876:133) continued, . . seven or eight bears were killed by our whole party . . . from the little black to the mammoth grizzly." Indians told Hoffman (1877:97) that the only place the black bear was encountered was in the Black Hills and in the Big- horn country. Bailey (1888:432) men- tioned that three kinds of bears (black, brown, and silvertip) "were said to oc- cur" in the Hills.
On 2 September 1968, a 160 pound ! domestic ewe, with its throat slashed and Hesh eaten from its back, shoulders, and hindquarters, was found in Swede Gulch, on the Philip Wolfe ranch, 2 mi N and 5 mi W Rochford, Pennington County (Rapid City Journal, 3 September 1968, p. 2). On the stream bank beside the ewe was the distinctive print of a bear. In November of the same year a 300- pound adult male black bear was shot west of Rochford, between Black Fox Campground and Crook's Tower.
Mr. Raymond Nilles, Forest Ranger of the Sundance District, told me of find- ing huge, deep claw marks on a tree along Hershey Creek in the northern Hills in 1966. The claw marks began about six feet from the base of the tree and extended to the ground; Mr. Nilles and others credit these markings to the activity of a bear. Other recent reliable sightings of black bear in the Black Hills are as follow: Big Spearfish Can- yon; along Cold Creek, in Lawrence County; near Redfern; in Beaver Creek Valley, in Pennington County; and at Camp Mallo in Weston County. It should be noted that captive black bears are tourist attractions in the Black Hills region, and some of these may occa- sionally escape.
Specimen examined ( 1 ) . — SOUTH DA- KOTA: Pennington Cotintij: west of Rochford, 1 (mounted and on display at Wall Drug Store, Wall, South Dakota).
Ursus aretos horribilis Ord
Grizzly Bear
C7r,v(/.v hornhilis Ord, 1815, in Guthrie, a new geograpliical, historical, and commercial grammar . . ., ed. 2, 2:291, 299 (type lo- cality, north side of Missouri River, near moutli of Wolf Creek, Roosevelt Co., Mon- tana ) .
Ursus aretos horribilis — Erdbrink, 1953, A re- vie\\' of fossil and recent bears of the Old World. . . . Jan de Lange, p. 339, 30 March.
Ursus rogersi bisonophagits Merriam, 1918, N. Amer. Fauna, 41:66, 9 February (type lo- cality. Bear Lodge Mountains, Sundance National Forest, Black Hills, Crook Co., Wyoming ) .
In discussing the Black Hills in 1851, a trapper indicated that "Grizzly bears are found there sometimes in bands like buffalo. . . ." (Culbertson, 1952:90). Although not quite as enthusiastic, Grin- nell (1875:81), Dodge (1876:132-1.33), and Bailey (1888:432) each alluded to the presence of large numbers of grizzly bears in the Black Hills in the late 1800's. The four specimens examined all were taken in the study area between 1855 and 1887. On 7 August 1874, General Custer and Colonel Ludlow shot a well- scarred old male near Nahant (about three miles south of the present town of Dumont, Lawrence County, South Da- kota); soon after, other members of the expedition killed a cinnamon-colored grizzly in the same area ( Winchell, 1875: 50; O'Hara, 1929:251-252). The male was a deep, glossy black, except for a sprinkling of dark gray hairs on the head, lower parts of the shoulders, and thighs (Grinnell, loc. cit.). Two Indian scouts killed an old female and two cubs a few days later much farther eastward (along Boxelder Creek, Lawrence Co., South Dakota). "The cubs were about half grown, and with the mother, were of a yellowish clay color. The inner half of each hair was deep black, but the outer extremity was a bright reddish- yellow. This gave them a curious mottled appearance and induced many of those who saw them to consider them a differ- ent species from the one killed by Gen- eral Custer" (Grinnell, loc. cit.).
128
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
The foregoing account points out the extreme amount of individual variation found among this species, and the taxo- nomic dilemma thus implied. At the moment, there are about 96 named kinds of grizzly bears, most of which can be attributed to the work of C. Hart Mer- riam. iMerriam (1918:66) described Ur- siis rogersi hisonophagus from the Bear Lodge Mountains, and at the same time (1918:93) included the Black Hills within the range of Ursus absarokos. I follow Erdbrink (1953:339), who re- garded the brown bear of the Palearctic and Nearctic regions as being conspe- cific and therefore employed the name Ursus arctos horrihiJis. Professor E. Ray- mond Hall, of The Universitv of Kansas, currently is reviewing the taxonomic status of the grizzly bear. The following selected cranial measurements of grizzlys from the Hills were taken by Professor Hall, and are the average (extremes) of three females and a male (type of U. r. hisonophagus) , all fairly young animals: occipitonasal length, 281.3 (255.0-322.0); palatal length, 166.5 (157.0-180.0); inter- orbital breadth, 70.5 (63.2-79.4); length of P4-M2, 72.2 (68.0-76.2); length of M2, 36.5 (33.7-38.4); breadth of M2, 18.1 (17.2-19.7).
Green (1961:147, 1962:3) recently lo- cated a partially fossilized arthritic skele- ton of a grizzly in a cave near Hot Springs, Fall River Co., South Dakota. It should be noted that all reports of bears in the Black Hills region since the turn of the century have been credited to U. americanus rather than U. arctos; evidently, even though rare in occurence, the former has replaced the latter in recent historic time.
Specimens examined ( 4 ) . — WYOMING : Crook County: Bear Lodge Mountains, 1 (USNM); Sundance National Forest, 3
(USNM).
Family PROCYONIDAE— Raccoons
Only one procyonid species, the rac- coon, is distributed in the Black Hills. The raccoon is an economically impor- tant mammal. Its pelt is marketable and
its flesh eaten by humans; it provides much recreation as a hunted fur-bearer, and accounts for substantial loss of cer- tain crops to farmers annually. The spe- cies probably is commoner in the region now than in the recent past.
Procyon lotor hirtus
Nelson and Goldman
Raccoon
Procyon lotor hirtus Nelson and Goldsmith, 1930, Jour. Mamm., 11:455, 11 November (type locality. Elk River, Sherburne Co., Minnesota ) .
Procyon lotor is common and wide- spread in suitable habitat throughout North America. Although few actual specimens have been taken, the species is common in the Black Hills. Usually the raccoon is found along streams, around lakes and marshes, and is a fre- quent nocturnal visitor to area camp- grounds.
Procyon lotor evidently is a recent addition to the Black Hills mammalian fauna. Hayden (1859:706, 1862:143) noted that this species was seldom seen beyond the frontier in the mid-nineteenth century, and indicated that, although a few had been obser\ed in the White River Valley, the raccoon seldom passed up the Missouri River above latitude 42°. Grinnell (1875) and Dodge (1876) did not include the raccoon in their respec- tive lists of mammals encountered by early expeditions to the Black Hills re- gion.
A careful search of the mud banks of almost any riparian community in the Hills will reveal the presence of tracks of P. lotor. While camping in Little Spearfish Ganyon in early August 1967, my cooking supplies were raided each night by a large male of this species. A huge steel trap, anchored by a heavy chain and baited with carcass of a chip- munk, finally captured the raccoon, but it escaped, dragging the trap behind. One week later a local fisherman found the animal, with the trap and chain still attached, drowned in Little Spearfish Greek. An adult Procyon and four ju-
TURNER: MAMMALS OF THE BLACK HILLS
129
\xMiilcs were observed in the area of Uoughlock Falls on 7 August of the same \ear. Another indi\idual was a nightly visitor at the Boxelder Creek Camp- ground, just \\ est of Nemo, in late June 1967. Specimens of this species obtained in the vicinity of Hill City were taken along a small tributary in a spruce can- yon.
Specimens examined (4). — SOUTH DA- KOTA: Lawrence County: 2 mi S, 10 nii W Lead, 5800 ft, 1. Pennington County: 4 mi SE Hill City, 5300 ft, 1 (UMMZ); S Hill City, 1 (U\LMZ). Custer County: Wind Cave Na- tional Park 1 (WCNP).
Additional records.— SOUTH DAKOTA: Custer County: Custer State Park (USES card files). WYONHNG: Crook County: New Haven (Goldman, 1950:38).
Family MUSTELIDAE— Weasels, Skunks, and Allies
Three genera and six species of weasel-like mammals inhabit the Black Hills; the occurrence of four additional genera ( five species ) in the general area remains uncertain. Members of this family mainly are terrestrial, but two (the badger and black-footed ferret) are semifossorial in habit, and one (the mink) is equally at home on land or in water. Breeding female mustelids gener- ally exhibit induced ovulation, delayed implantation, and are monestrous. Both sexes possess anal scent glands, of which those of weasels, mink and skunks are the most noisome to humans.
Mustela erminea muricus (Bangs)
Ermine
Putorius (Arctogale) muricus Bangs, 1899, Proc. New England Zool. Club, 1:71, 31 July (type locality. Echo, 7500 ft, El Do- rado Co., California).
Mustela erminea murica — Hall, 1945, Univ. of Kansas Publ, Mus. Nat. Hist., 25:84, 27 February.
Mustela erminea muricus is distrib- uted throughout the northwestern quar- ter of the United States; in the Black Hills it is most common in lower spruce- aspen parklands, streamside coniferous forest, and grassy to semi-swampy wood- lands, usually at altitudes above 5000
feet. This small weasel is rather un- common in collections due to its seclusive habits and is known only from Penning- ton and Custer counties in the South Dakotan portion of the Hills, but from both counties of the Wyoming section. Hall (1951:164-165) first reported this species from the study area.
Ecological data concerning the er- mine in the Black Hills are rather meager. During the winter months of 1945-1946, J. A. King captured five indi- viduals (two males and tliree females) in the central Hills in box traps. The traps were placed in aspen groves and along marshes bordering small tribu- taries that flowed through canyons domi- nated by spruce. One female that was kept as a pet by King, carried four em- bryos when she died on 30 March 1946. Two females were obtained in mid-July of 1965 on a huge dirt slide, overgrown with grass and small aspen, situated be- tween two high, rocky outcrops east of Four Corners, Wyoming. One had raised a fitter shortly before capture; three old placental scars were present and mam- mary tissue was receding. The other was a subadult, possibly a young of the first female. Coats (1945:10) captured 16 ermine one winter in the vicinity of Hill City.
Individuals taken in the warmer months (June and July) were in brown summer pelage and those obtained in colder months (November to January) were in white winter pelage. Three er- mines that died in March were molting from winter to summer pelage; two of these had been kept as pets by King since mid-January, and were in the 21st day (UMMZ 90189) and 37th day (UMMZ 90190) of molt, respectively, at time of death.
Specimens examined (12). — SOUTH DA- KOTA: Pennington County: Palmer Gulch, 3 mi SE Hill City, 5300 ft, 2 (UMMZ); Pflan- der's Ranch, 3 mi SSE Hill City, 5300 ft, 1 (UMMZ); Nelson's Place, 4 mi SE Hill City, 5300 ft, 1 (UMMZ); Spring Creek, south of Rapid Citv, 2 (MHM); Spring Creek, 2 mi W Ore%ille, 5500 ft, 1 (UMMZ). Custer County: 0.5 mi E Sylvan Lake, 6250 ft, 1 (UMMZ).
WYOMING: Crook County: 3 mi NW
130
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Sundance, 5900 ft, 1. Weston County: 4 mi E Four Corners, 6000 ft, 2.
UNSPECIFIED LOCALITY: Black Hills, 1 (SDSU).
Mustek frenata alleni (Merriam) Long-tailed Weasel
Ptitorius alleni Merriam, 1896, N. Amer. Fauna, 11:24, 30 June (type locality, Black Hills, Custer, Custer Co., South Dakota).
Mustela frenata alleni — Hall, 1936, Carnegie Inst. Washington Publ., 473:106, 20 Novem- ber.
The long-tailed weasel occurs throughout the United States. In the Black Hills it inhabits brushy areas, woodlands, and rock piles, usually above 5000 feet and near a source of permanent water. The species e\'idently is rather uncommon in the Hills, being presently kno\\aT[ only from Pennington, Custer, and Crook counties. This mustelid was first recorded from the study area by Vernon Bailey (1888:433).
As presently understood, Mustela frenata alleni is confined to the Black Hills of South Dakota and Wyoming. Hall (1951:276) recorded alleni from Mitchell, Scott's Bluff Co., Nebraska, but Jones (1964:270) subsequently assigned these specimens to the subspecies longi- caucla. Specimens from northwestern Nebraska, and one from Rapid City, Pennington Co., South Dakota, undoubt- edly represent intergrades between alleni and longicauda.
Mustela frenata alleni difters from M. f. longicauda, which occurs to the north, east and south of the Black Hills, in possessing smaller cranial and external dimensions. Mustela frenata nevadensis, known from the west in Wyoming, aver- ages slightly larger in size and is a darker brown in summer pelage than is M. f. alleni (Long, 1965:693).
Ecological data are essentially lack- ing for this species in the Black Hills. In November 1887, Vernon Bailey caught a long-tailed weasel in a trap baited with a prairie dog and set on a creek bank near Rapid City. During the summer of 1968, individuals were observed along stream drainages, brushy ravines, and
around headquarters buildings in Wind Cave National Park, but no specimens were obtained. J. A. King captured a male in an aspen thicket southeast of Hill City in January of 1946; this indi- vidual was retained in capti\ity until 9 April. King noted that molt from white winter pelage to brown summer pelage commenced about 4 February; thus, when the male (UMMZ 90209) died, it was in the 64th day of pelage replace- ment and long white guard hairs were still present over shoulders and neck. Bailey ( loc. cit. ) noted that the Novem- ber-taken individual ". . . is just chang- ing from the brown coat to the white, and is a little more than half white. The brown hairs come out very easily; the white hairs are firm." He also witnessed a few tracks along creeks and ventured that long-tailed weasels were quite com- mon in the general area.
Specimens examined (13).— SOUTH DA- KOTA: Pennington County: Hill City, 1 (AMNH); Palmer Gulch, 3 mi SE Hill City, 1 (UMMZ); 20 mi N Elk Mountain, 6000 ft, 1 (USNM). Custer County: Palmer Gulch, 8 mi SE Hill City, 5300 ft, 4 (FMNH); Bull Springs, 2 mi N, 9 mi W Custer, 1 (UMMZ); Custer, 3 (2 AMNH, 1 USNM).
WYOMING: Crook County: Bear Lodge Mountains, 3 mi N Sundance, 5500 ft, 1 (USNM); Sundance, 1 (USNM).
Mustela nigripes (Audubon and
Bachman ) Black-footed Ferret
Putorius nigripes Audubon and Bachman, 1851, The viviparous quadrupeds of North Amer- ica, 2:297 (type locality. Ft. Laramie, Goshen Co., Wyoming, according to Hay- den, Trans. Amer. Philos. Soc, 12 (n. s. ): 138, 1863).
Mustela nigripes— MiWer, 1912, Bull. U. S. Nat. Mus., 79:102, 31 December.
Presently on the list of endangered species, the black-footed ferret is among the rarest of North American carnivores. This mustelid is characteristic of short and mid-grass prairies, and its geographic range is generally coincident with that of the sciurid genus Cijnomys. Although ferrets utilize alternate sources of food (Cahalane, 1954:423; Hillman, 1968:6; Henderson et al, 1969:22-24), prairie
TURNER: MAMMALS OF THE BLACK HILLS
131
dogs are its primary prc\^ and burrow systems of the latter species are inhabited by ferrets.
Mustela niiiripes currently seems to be holding its own in western South Dakota, but changing land use and man's effort to control prairie dog populations through destruction of thousands of dog towns have adversely affected the distri- bution and abundance of black-footed ferrets throughout its range. For ex- ample, widespread use of compound 1080 to eradicate Cijnomys is a potential threat to the survival of ferrets. Hillman (1968:11-12) has shown that consump- tion of 1080-poisoned prairie dogs by black-footed ferrets can cause fatal sec- ondary poisoning.
Henderson et al (1969:31-37) re- cently summarized the distributional status of M. nigripes in South Dakota. His census was based on questionnaires gi\'en to land owners, on a survey of published literature and unpublished records such as those located in the U. S. Biological Survey files, and on examina- tion of specimens housed in various mu- seums. Although the species was not un- common on the prairie along the eastern periphery of the Hills, only a few records are directly applicable to the Black Hills proper.
In 1903-1904, James P. Campbell ob- served a ferret in a prairie dog town in Gillette Canvon, northeast of Elk Moun- tain, Custer County. Another individual was observed running along a road lo- cated in T. 2 S, R. 5 E, 5800-6000 feet, Custer County, by James Clark in July 1965; this site may be within the foot- hill region of Custer State Park, where habitat would be suitable for ferrets.
Troy C. Beach observed a M. nigripes in a prairie dog town in Wind Cave National Park in the spring of 1922. Upon completion of filming "The Van- ishing Prairie," Walt Disney Productions, Inc., released two males and one female in the National Park on 30 December 1953 (Cahalane, 19.54:424; Garst, 1954: 594). These were three of five ferrets (an adult and four young) trapped in a
dog town 2 mi W Midland, Haakon Co., South Dakota, by Mr. George Barnes early in August 1953 (Garst, loc. cit.). Subsequently, a Park Naturalist noted a ferret in the vicinity of Norbeck Dam on 27 August 1956. The last actual sight- ings of this species in the Park were in 1957 (Wind Cave National Park files). When excavating prairie dog burrows, ferrets form a characteristic trench-like structure from three to five inches in width and from one to nine feet in length (Henderson et al, 1969:16). In 1969, the diagnostic trench of a black-footed ferret was noted in a dog town in Wind Cave National Park. Restocking of M. nigripes in the National Park, and in other sanctuaries where prairie dogs thrive, such as Custer State Park and Devils Tower National Monument, could insure the survival of this interesting mammal.
Little is known concerning the repro- duction of the black-footed ferret but young evidently are born in spring and first emerge above ground early in July, at which time family groups are most frequently observed in western South Dakota.
Specimens examined (4). — SOUTH DA- KOTA: Pennington County: Spring Creek, south of Rapid City, 2 (MHM).
WYOMING: Weston County: Newcastle, 2 (USNM).
Additional records. — (see text and Hender- son et a/.: 31-37, Appendix H).
Mustela vison letifera Hollister Mink
Mustela vison letifera Hollister, 1913, Proc. U. S. Nat. Mus., 44:475, 18 April (type locality, Elk River, Sherburne Co., Minne- sota).
Mink live along streams, edges of ponds, marshes, and lakes in all but the southwestern part of the United States. Uncommon in collections due to its valu- able pelt, Mustela vison is fairly abun- dant along water courses throughout the Black Hills. For example, "numerous mink" are trapped near Sturgis annually (South Dakota Conservation Digest, 23: 12, March 1956).
132
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Grinnell (1875:80) saw no living specimens while in the Hills, but ob- served signs of mink along streams and saw a skin that the owner said had been taken "in the Black Hills." Bailey (1888: 433) noted the presence of this species in the Hills in November 1887.
A lactating female was taken along the edge of a beaver pond south of Tinton on 16 July 1961 by W. C. Stanley; to my knowledge, no additional data are available for this species from the Black Hills, save for the specimens listed below.
Specimens examined (4). — SOUTH DA- KOTA: Lawrence County: 2 nii S Tinton, 6100 ft, 1. Pennington County: Rapid City, 1; Martindale Ranch, 0.5 mi E Rapid City, 1 (SDMT); Spring Creek, south of Rapid City, 1 (MHM).
Taxidea taxus taxus (Schreber) Badger
Ursus taxus Schreber, 1778, Die Saugthiere . . ., description on pp. 520-521 (type locality, Labrador and Hudson Bay, restricted south- west of Hudson Bay by Miller and Kellogg, Bull. U. S. Nat. Mus., 205:747, 3 March 1955).
Taxidea taxus — Rhoads, 1894, Amer. Nat. 28: 524, June.
Taxidea taxus is a relatively common inhabitant of the Northern Great Plains surrounding the Black Hills, where it occurs throughout the Red Valley "race- track," Wind Cave National Park, and other areas where fringes of prairie im- pinge upon the foothills. Being a rapid and powerful digger, the badger pro- vides reliable evidence of its presence by its characteristic excavations and den sites, usually located on gently sloping hillsides.
The genus Taxidea, with but one Recent species, is distributed in centi'al and western North America. Some years ago, Viola S. Schantz began a revision, naming 10 new subspecies over a period of several years, but a final report was not issued; Charles A. Long currently is analyzing geographical variation of Taxi- dea taxus. Schantz (1946:81) described T. t. dacotensis from Folsom, Custer Co., South Dakota, just east of the Black Hills, and included specimens from the Hills
under that name. Because of the great individual variation in the species and because an adequate number of speci- mens for analysis is lacking, I tentatively follow Hall ( 1936) in assigning the Black Hills specimens to Taxidea taxus taxus.
The badger is rather uncommon in the Black Hills proper. Cahalane (1948: 249; 1951:209) reported that a pair was released about 1940 in Wind Cave Na- tional Park, where tliriving prairie dog towns offer an ample food supply, and that this pair produced at least one litter of young. Koford (1958:31) noted that several prairie dog tunnel systems in Shirttail Canyon had been excavated by badgers in 1955. In the early morning of 9 July 1968, I observed an adult female and two pups digging out a Cynomys burrow on Bison Flats, in the National Park; a Ranger-NaturaHst at the Park observed a female badger carrying a young pup in her mouth near that prairie dog town on 20 May 1957 (Wind Cave National Park files). An adult T. taxus was observed along the road 6 mi NW Spearfish on 5 August 1969 eating a road- , killed Sciurus niger.
Bounty claims were paid in 1962 on 37 badgers taken in the counties that comprise the South Dakotan portion of the Black Hills (records from South Da- kota Dept. Game, Fish and Parks ) . One indi\'idual trapped in the winter of 1964- 65 southwest of Hill City was taken at an elevation in excess of 6000 feet.
Specimens examined (6). — SOUTH DA- KOTA: Meade County: 5.6 mi N Piedmont, 1 (SDMT). Pennington County: near Rapid City, 1 (FMNH); Spring Creek, south of Rapid City, 1 (MHM); 4 mi S, 14 mi W Hill City, 1. Custer County: Dewey, 2 (USNM).
Mephitis mephitis hudsonica
Richardson
Striped Skunk
Mephitis americana var. hudsonica Richardson, 1829, Fauna Boreali-Americana, 1:55 (type locality, plains of Saskatchewan, Canada).
Mephitis mephitis Jiudsonica — Hall, 1934, Univ. California Publ. Zool., 40:368, 5 November.
The shiped skunk occurs throughout the United States. In the Black Hills, it
TURNER: MAMMALS OF THE BLACK HILLS
133
ranges from the lo\^or elevations of the sunoimding plains, through the foothill ti-ansition zone, up to altitudes of about 6200 feet. The species is at home in a variety' of habitats, but prefers forest borders, wooded ravines, rocky outcrops, and brushy agricultural fields with asso- ciated fencerows. It is no sti'anger to the habitations of man, and although represented in museum collections by only a few specimens, the species is com- mon throughout the Black Hills region. Grinnell (1875:80) first noted the pres- ence of this mustelid in the Hills.
Striped skunks often were observed along roadsides and in the vicinity of campgrounds. Residents of Nemo, Law- ence County, indicated that more than 70 individuals had been killed in the community^ dump in 1966-1967; an adult female and an immature female were obtained there on 30 June 1967. Bailey (1888:433) captured an individual near Rapid City in a trap baited with the car- cass of a prairie dog. Most remaining specimens taken in the Black Hills have been either killed on roads, or represent skeletal material found in caves and rockv crevices.
A juvenile was captured on 8 August 1894 at Custer by W. W. Granger. Another immature Mephitis that was ob- viously sick was captured along a road northwest of Minnekahta on 16 June 1967. This indi\'idual offered no resist- ance; possibly it had been poisoned or was rabid. A female taken near Nemo on 30 June had five placental scars, two in the right uterine horn, and three in the left.
Specimens examined (30). — SOUTH DA- KOTA: Lawrence County: Nemo, 4700 ft, 2. Pennington County: near Rapid City, 8 (FMNH); Moon, 22 mi W Hill City, 6200 ft, 1; Ditch Creek Valley, 14 mi W Hill City, 6400 ft, 1; Hill City, 4975 ft, 2; Spring Creek, 3 mi E Hill City, 1 (UMMZ); Spring Creek, south of Rapid City, 1 ( MHM ) ; 20 mi N Elk Mountain, 1 (USNM). Custer County: Custer, 3 (1 USNM, 2 AMNH); south of Otis, 1 (UMMZ); Wind Cave National Park, 1 (WCNP); Elk Mountain, 1 (USNM); Dewey, 3 (USNM). Fall River County: 1.75 mi N, 4.5
mi W Minnekahta, 3800 ft, 1; 5.5 mi E Minne- kahta, 4000 ft, 2.
WYOMING: Crook County: 3.5 mi NW Sundance, 5000 ft, 1.
Additional record— SOUTH DAKOTA: Fall River County: Minnekahta (USES card files).
Family FE LI DAE— Cats
Of the three felids that occur in the Black Hills, bobcats are by far the most common; mountain lions are less fre- quent, and lynx are but rare visitors to the region in recent years.
Felis concolor hippolestes Merriam Mountain Lion
Felis hippolestes Merriam, 1897, Proc. Biol. Soc. Washington, 11:219, 5 July (type locality, Wind River Mountains, near head Big Wind River, Fremont Co., Wyoming).
Felis concolor hippolestes — Nelson and Cold- man, 1929, Jour. Mamm., 10:347, 11 No- vember.
Previously, Felis concolor occurred through much of North America, but it has been extirpated in many parts of its range and now occurs primarily in re- gions poorly populated by humans. The mountain lion feeds principally on deer and occasionally upon livestock. Due to its role as a large predator, this species has been persecuted to the point that even within its present range, it is now rare and should be protected. Felis con- color is native to the Black Hills, although it is uncommon and inhabits the more remote and rugged areas.
Members of the Custer Expedition of 1874 saw but one "panther," near the head of Castle Creek (O'Hara, 1929: 252). Grinnell (1875:79), who accom- panied Custer, believed that these felids were quite numerous and saw indications of their presence on several occasions: "I . . . once found the partially de- voured remains of a deer thad had just been left by one of these animals." Mem- bers of the Newton- Jenney Survey ob- served two or three mountain lions but none was killed (Dodge 1876:133). A female mountain lion and two young were shot in November 1887, just after the adult had killed a large mule deer buck (Bailey, 1888:431) and Merritt
134
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Gary obtained a female near the head of Bear Creek in the Bear Lodge Moun- tains in 1890.
Earl Bedell shot a lion near the head of Stockade Beaver Creek, Weston Co., Wyoming, in 1930 (Mann, 1959:3) and a female F. concolor was killed early in December 1931 near the Wyoming bor- der (Yomig, 1946b: 34). The latter indi- vidual was known in the region for the previous two years and was believed to have raised at least one litter in this period. In a report written to the U. S. Biological Survey in 1932, Louis Knowles indicated that these were the only moun- tain lions taken in the Hills in the pre- vious 25 years.
The most recent specimen taken in the Black Hills to my knowledge was shot by Ted Mann on 16 December 1958 (Mann, 1959:5). He located fresh tracks in the snow on Elk Mountain, in Custer County. In the company of two neigh- bors and three dogs, Mann set out in pursuit of the lion, which had killed a doe and was eating on the hindquarters when the dogs jumped it. The lion, a male that weighed 140 pounds and meas- ured about seven feet in length, finally was treed and shot along Upper Dugout Creek, about two miles south of the Steam's Ranch.
In 1965, Fred A. Fichtner, District Ranger from Newcastle, Wyoming, re- ported that there had been several re- cent reports by ranchers of mountain lion tracks between Custer and Elk Mountain, South Dakota; "Plaster casts were made of one set, and positively identified" (letter to J. Knox Jones, Jr.). Ranger A. W. Jones, Spearfish District, noted the recent presence (1968) of a mountain lion in the vicinity of Big Crow Peak (pors. com.). Park Ranger Fred Devenport (pers. com.) indicated that two F. concolor were observed just in- side of the west boundaiy of Wind Cave National Park on 10 April 1964 by Dr. Ralph Hubbard and an Indian com- panion, of Wounded Knee, South Da- kota. Ranger-Naturalists reported see- ing mountain lions in the Park on two
different occasions in 1965, and on 2 April of that year, fresh lion tracks were identified in soft mud next to the pump house. A lion also was observed about this time on the ranch of Mr. Shirlev McClure, southwest of Hot Springs.
Specimen examined (1). — WYOMING: Crook County: head Bear Creek, Bear Lodge Mountains, 1 (USNM).
Lynx canadensis canadensis Kerr Lynx
Lynx canadensis Kerr, 1792, The animal king- dom . . ., 1:L57 (type locality restricted to Pro\ince of Quebec, Canada, by Miller and Kellogg, Bull. U. S. Nat. Mus., 205: 777, 3 March 1955).
Lynx canadensis is typical of the heavily forested boreal regions of North America, but formerly occurred spar- ingly in suitable habitat in the Northern Creat Plains region. Grinnell (1875:79) and Dodge (1876:12.3) both indicated that this species previously inhabited the Black Hills, and there have been several recent reports of lynx in the area. In 1944, a Ivnx was killed in Meade Countv and two more were taken in Pennington Countv (South Dakota Conservation Di- gest, 12:15, May 1945). A. H. Richard- son, State Game Biologist, wrote (pers. com.) that in the last 10 years he has personal!)' examined two specimens shot in the northern and western Black Hills.
I know of but three other individuals captured in South Dakota. A lynx was taken in Granger's Woods along the Missouri River, above Sioux Citv, in January 1875 (Brackett, 1881:413). A male (USNM 246.548) was obtained by J. N. Martin at Bullhead, Corson County, on 6 October 1925, and another was shot by Mr. Leroy Johnson near Marindahl, Yankton County, on 6 May 1962 (Lee, 1962:21). Lynx canadensis presently in- habits eastern Montana (Hoffman and Pattic, 1968:. 53), and has been taken south of the Black Hills in the Laramie Mountains, Albany County, Wyoming (Long, 1965:708) and in northern Ne- braska (Jones, 1964:304).
TURNER: MAMMALS OF THE BLACK HILLS
135
Lynx rufus pallescens Merriam Bobcat
Lynx fascidtiLs pallescens Merriam, 1899, N. Amer. Fauna, 16:104, 28 October (type lo- cality, S side Mt. Adams, near Trout Lake, Skamania Co., Wasliington ) .
[Lynx rufa] pallescens — Elliott, 1901, Field Columb. Mus., Publ. 45, Zool. ser., 2:297, 6 March.
The bobcat is widely distributed in the Black Hills, being common in the foothills, in forested canyons, along wooded streams, and in areas of exposed rimrock. It remains abundant in the Hills region in spite of predator control measures. Bounties on 1311 bobcats were paid by those counties that com- prise the South Dakotan portion of the Black Hills from 1962 to 1968 (records from South Dakota Dept. Game, Fish and Parks). In 30 years as a Federal Trapper, Archie Howe of Custer killed more than 450 bobcats in the Hills (Popowski, 1952:6).
Although he saw no specimens, Grin- nell (1875:79) indicated that this felid was common in the Hills region. Bailey (1888:432) noted the presence of L. rufus near Deadwood in October 1887, and collected one individual near Custer in July 1888.
Evidently, the bobcat is an efficient predator on large game mammals. Young (1958:2.3) reported that "A large buck antelope was no match for a bobcat which killed it one winter day of early 1930 on the Wind Cave Game Preserve [Wind Cave National Park] in South Dakota." On 5 April 1954, George Barnes shot a 27-pound male that had just killed a yearling white-tailed buck near Wabash Spring, 5 mi N of Custer {op. c?Y.:84). However, small game also is included in the diet of this species. Stomachs of two females trapped in Wind Cave National Park by Archie Howe in early June 1924 contained only the remains of rabbits (U. S. Biological Survey files ) .
In January 1946, a farmer found a dead beaver, with a trap attached, float- ing under the ice on a stream in Palmer
Gulch. He placed the carcass on the bank, set the trap, and captured three bobcats within a week (J. A. King, field notes). Stebler (1939:389) obtained a specimen of L. rufus along Grace Cool- idge Creek in Custer State Park in Au- gust 1934. The stream was bordered with bur oak, quaking aspen, paper birch, and other deciduous trees and shrubs.
Both of the females taken in Wind Cave National Park in early June 1924 were lactating. On 17 August 1967, I shot a large female and two three-quarter grown kittens at dusk as they were leav- ing a den at the base of a limestone out- crop just southeast of the National Park. Thus, females of this species probably give birth to young in late March or early April in the Black Hills.
Specimens examined (38). — SOUTH DA- KOTA: Meade County: 6 mi E Piedmont, 1 (SDSU). Pennington County: near Rapid City, 1 ( FMNH ) ; Spring Creek, south of Rapid City, 1 (MHM); Palmer Gulch, SE Hill City, 3 (UMMZ); 4 mi S, 14 mi W Hill City, 1; un- specified locality, 1 (NRW). Custer County: S Otis, 1 (UMMZ); Custer, 3 (USNM); French Creek, near Custer, 5 (USNM); Bowman Ridge, near Custer, 1 (USNM); Beaver Creek, south of Custer, 1 (USNM); Mayo, south of Custer, 1 (USNM); Streeter Ranch, 1.5 mi S, 5 mi E Wind Cave, 3550 ft, 3; Elk Mountain, 1 (USNM); Dewey, 6 (USNM).
WYOMING: Crook County: Otto, 1 (AMNH); Leggot Canyon, Sundance, 1 (USNM); Person Canyon, Sundance, 1 (USNM); Sand Creek Canyon, 1 (USNM). Weston County: Newcastle, 1 (USNM).
UNSPECIFIED LOCALITY: Black Hills, 3 (2 USNM, 1 WCNP).
Additional records ( see text also ) . — SOUTH DAKOTA: Custer County: Bull Springs, 14 mi NW Custer (Lamster, 1943:1). County unspecified: Calumet Draw (South Dakota Conservation Digest, 17:5, June 1950).
ORDER ARTIODACTYLA—
Even-toed Ungulates
Artiodactyls in the Black Hills belong to the suborder Ruminantia, and are rep- resented by three families with five gen- era and six species. A seventh species, the mountain goat, was introduced from western montane areas. An additional species, the fallow deer, has been intro-
136
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
duced from the Old World, but is not well established in the Hills as yet. Four of the native species, wapiti, pronghorn, bison, and mountain sheep, previously were extirpated or drastically reduced in numbers, and subsequently have been reintroduced from other areas. Artio- dactyls comprise a varied assemblage of hooved and horned, or antlered, mam- mals that were economically important to early explorers and pioneers. The pres- ent-day game species still provide rec- reation for outdoorsmen and much eco- logical information in the following accounts is drawn from conservation and game management literature.
Family CERVIDAE — Elk and Deer
Cervids are mainly browsers. Antlers are present at least in males and are shed annually. This family is represented in the Black Hills by two genera and three species.
Cervus canadensis canadensis Erxleben
Wapiti
[Cervus elaphtis] canadensis Erxleben, 1777, Systema regni animalis . . ., p. 305 (type locality, eastern Canada, probal:)ly the vi- cinity of Montreal according to V. Bailey, Proc. Biol. Soc. Washington, 48:187, 15 November 1935).
The wapiti, or American elk formerly occurred throughout most of temperate North America; it has been reintroduced in many areas where it previously had been extirpated by man. The subspecies C. c. canadensis was native to the Black Hills, but, once it became extinct, the area was restocked with C. c. nelsoni from Wyoming. Cervus canadensis may be conspecific with the earlier-named red deer, Cervus elaphus, of Eurasia (McCullough, 1969), but I retain cana- densis pending definitive study of their relationships.
The Custer Expedition of 1874 saw only a few elk, but several were killed along Boxelder Creek near Nemo on 11 August (Winchell, 1875:5.3; O'Hara, 1929 : 252 ) . Grinnell ( 1875 : 83 ) , who ac- companied General Custer, gave the fol-
lowing account that partially indicated the prior abundance of wapiti in the Hills: "On Elkhorn Prairie [Reynolds Prairie] we came upon a collection of liorns gathered together by the Indians. Three lodge-poles had been set up in the ground so as to form a tripod, and supported by these were a pile of [wa- piti] horns 8 to 10 feet high. The horns had been shed, and had apparently been collected from the surrounding prairie [isolated prairie in the middle of the Black Hills] and heaped up here by the Indians." Dodge (1876:132), describing a hunt along the northern rim of the Black Hills, noted that a hunter "fol- lowed a large elk trail into a little valley" that was densely vegetated and that at the report of his gun, "the whole valley was alive with rushing animals; at least a hundred, crashing through the brush, disappeared up the mountain side." Cer- vus canadensis was present in the lime- stone countrv of the Black Hills, Bear Lodge Mountains, Sundance Mountains, and along Stockard Beaver and Inyan- kara Creeks, Weston Countv, Wvoming, in 1885 (Murie, 1951:42),' and Bailey (1888:434) noted that some were "said to occur still" in the Black Hills in 1887.
Near the turn of the century, J. A. Allen (1895a: 263) indicated that the \\'apiti had been extinct in the Black Hills for se\'eral years. Thus, the original population of wapiti (C. c. canadensis) was mostly exterminated from the Hills by 1900, and initial restocking with C c. nelsoni from Wyoming commenced. About 65 were stocked at Custer State Park in 1912, but these escaped into the surrounding Hills and were replaced by an additional 25 in 1913; 50 more of these large cerxids were obtained from Mon- tana in 1916. Wind Ca\e National Park initiated a wapiti herd with 21 animals from Jackson Hole and Yellowstone Na- tional Park in 1914. Currently there are about 1200 wapiti in the State Park, and 450 in the National Park (Cahalane, 1948:253). The herds are controlled by drixing excess animals out of the confines of the parks into the surrounding Black
TURNER: MAMMALS OI-^ TIIK HLACK HILLS
137
Hills, or by issuing hunting permits each year.
PresentK, there are two herds, total- ing about 1000 elk, ranging through the Black Hills (Evans, 1966:9). A small "northern herd" traverses the limestone region along the South Dakota-Wyoming border, and a "southern herd" in habits the area adjacent to the National and State parks (A. H. Richardson, pers. com.). In the 1966-67 hunting season, 198 wapiti were killed by hunters in Custer State Park, 40 ( 19 bulls, 15 cows, and six calves) were taken from the "northern herd," and 40 (21 bulls, 13 cows, and six calves) were shot from the "southern herd" (Rapid City Journal, p. 9, 15 January 1967).
There is much variation in the antlers of this species in the Black Hills, and deformations are not uncommon; some antlers are much flattened near the ex- tremities, being somewhat palmate in nature. Wapiti retire to dense forests during daylight hours, descending onto the \'alley meadows in early evening. Bugling of males can be heard in the autumn during mating season, and caKes are born in late May or early June.
Specimens examined (4). — SOUTH DA- KOTA: Lawrence County: 2 mi S Tinton, 6100 ft, 2. Custer Countif: 1 mi NW Wind Cave, Wind Ca^■e National Park, 1 (WCNP).
WYOMING: Weston County: 0..5 mi E Buckhorn, 6150 ft, 1.
Odocoileus hemionus hemionus
(Rafinesque)
Mule Deer
Cervus hemionus Rafinesque, 1817, Anier. Monthly Mag., 1:436, October (type lo- cality, mouth of the Big Sioux River, South Dakota ) .
Odocoileus hemionus — Merriam, 1898, Proc. Biol. Soc. Washington, 12:100, 30 April.
The mule deer, or black-tailed deer, inhabits broken country, open plains, and brush or woods of western North America. An estimated 10 million mule deer inhabited the continent when Eu- ropean man arrived (Merwin, 1971:3.3). This species occurs throughout the Black Hills but is most abundant in the ra-
vines of foothills, and in the more remote and rugged sections, lioth the mule deer and white-tailed deer were an im- portant somce of food for early travelers and settlers of the region; accounts of Hayden (1856:79; 1859:706), Grinnell (1875:84), Dodge (1876:131), Bailey (1888:434), and J. A. Allen (1895a:283) indicated that Odocoileus hemionus was present in the Hills in considerable num- bers. However, market hunting and other factors essentially eliminated deer from most of South Dakota in the early 1900's (Seton, 1929b : .332 ); only 225 deer, mostly O. hemionus, were taken legally in South Dakota in 1920 (op. cit. -.250). A "buck law" and other legislation passed in the period between 1911 and 1925 enabled the populations of deer to in- crease in abundance once more by 1940 (Merwin, 1971:33). Currently, over- population and o\'er-utilization of avail- able range are the major problems in managing the ever increasing Black Hills deer herd.
The number of deer in the Hills in 1969 was estimated at 103,846—23 per- cent of which \\'ere O. hemionus (Rich- ardson, 1969b: 25; Richardson and Russel, 1970:3). Pellet group counts (based on a 365-day accumulation) conducted along 107 belt transects in 1970 indicated a 16 percent decrease in the Black Hills deer population over that of 1969. Northern Hills herd decreased 23.0 percent; cen- tral Hills herd decreased 14.6 percent, and southern Hills herd increased 5.3 percent (Thompson and Hausle, 1971:8). Mule deer are nati\'e to the Black Hills, but the species currently seems to be declining in abundance, occurring only in small pockets throughout the region, with greatest concentrations in the south- ern portion of the Hills. The ratio of white-tailed deer to mule deer varies from area to area. The herd in Wind Cave National Park and that of Fall River County are composed almost en- tirely of mule deer (F. Devenport, pers. com. ) , whereas in the western section of Custer County, there are equal numbers of each species. However, mule deer
138
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
predominate on the "West River Prairies" immediately adjacent to the Black Hills, with the white-tail being essentially a dweller of wooded rixer bottoms ( Bever, 1957:4); for example, 74 percent of the 12,910 deer taken by hunters in this area in 1968 were O. hemionus (South Da- kota Conservation Digest, 36:4, Decem- ber, 1969).
Projected kill of deer in the 3517 square miles of the South Dakotan por- tion of the Black Hills in 1968 was 4.02 deer per square mile and was composed of 31 percent white-tail does and 45 per- cent white-tailed bucks, 10 percent mule does and 14 percent mule bucks (Rose 1970, Tables 4-6).
South Dakota State Game Biologist A. H. Richardson (pers. com.) suggests that one of the main reasons for the de- crease of mule deer in the Hills is the preference of this species for an open, rolling type of habitat; in the past 20 to 40 years, ponderosa pine has enclosed many former environs of this type (Bever, 1959:4). In addition, white-tails seem to be the more aggressive of the two species in the Black Hills, forcing out the mule deer. Also, relative per- centage of mule deer killed in the Hills by hunters is greater than that of white- tailed deer. Differential response of the two species to harsh weather conditions suggests another possible explanation of the problem. For example, from 29 April to 4 May 1953, snow in excess of 44 inches fell in the northern Hills. Odo- coileiis vir^iniamis, in the higher por- tion of the Hills, migrated 10 to 25 miles down to the lower slopes and foothills during the winter for better browse and more equitable weather conditions. Mule deer did not migrate as readily, but simply shifted to more favorable expo- sures in nearbv steep canyon drainages (Berner, 1953b: 10; Harris, 1953:16; Hodgins, 1956:14).
Southwest slopes, from which the snows melt most rapidly, and tops of ridges, from which the snows are readily blown by w inds, are areas on which the deer concentrate in the winter. The vi-
cinity of Annie Creek, Lawrence County, serves as the main winter range for mule deer, whereas white-tailed deer over- winter on the Hepler Range of the north- ern foothills. Both species utilize the Aztec Hill, McVey Burn, Crow Peak, and Piedmont winter ranges. The criti- cal period for deer in the Black Hills is from January to April, and many cases of death due to malnutrition in winter have been recorded ( Gastler et at., 1951 : 356). The severe blizzard in the winter of 1948-1949 resulted in the deaths of 40 deer per square mile in the McVey Burn region (Berner, 1953a :4). Mule deer are especially susceptible to starvation and large herds in Spearlish Canyon must be fed high protein concentrate each winter due to overuse of the winter- ing grounds (Harris, 1952a:7), 1953:14; Brady, 1955:13; Leopold, 1956:4). Such aggregations in restricted areas further complicate the problems. A survey of browse in April and May 1969 indicated that in 2633 square miles of wintering range in the Hills, 92 square miles were over-utilized; 208 square miles were properly utilized, and 2333 square miles were under-utilized (Richardson, 1969: 3). Over-utilization occurred on the McVey Burn, Crow Peak, and Piedmont ranges, and in the area of mountain mahogany west of Custer.
Rut extends from early November to mid-December. Fawns, usually twins, are born in the Black Hills in May and June. Deer lice (Lepoptena depressa) parasitize deer of the Wind Ca\e Na- tional Park herd externally (Wind Ca^'e National Park files).
Specimens examined (24). — SOUTH DA- KOTA: Lawrence County: 1 mi S, 4 mi W Spearfish, 1. Pennington Cotintt/: Rapid City, 1 (USD); Beaver Creek Vallev, 4 mi N, 10.5 mi W Deerfield, 6400 ft, 1; 3 mi N, 4 mi E Hill City, 1. Custer Cotinti/: Pleasant Valley Ranch, 6 mi SW Custer, 2'(AMNH); Camp- hell's Ranch, Elk Mountain, 4800 ft, 1 (USNM); Redhird Canvon, Elk Mountain, 6000 ft, 1 (USNM); Elk Mountain, 5 (USNM). Fall River Countt/: 1 mi N, 4 mi E Edgeniont, 2600 ft, 1.
WYOMINC: Crook County: Bear Lodge Mountains [not 9524 (=U. S. Biological Sur-
TURNER: MAMMALS OF THE BLACK HILLS
139
vey skull number) feet as noted hv Long, 1965:713—7242 feet is the hij^hest ele\ ation in the Blaek Hills], 2 (USNM). Weston County: 3 mi E Newcastle, 1; unspecified lo- cality, 7.
Odocoileus virginianus dacotensis
Goldman and Kellogg
White-tailed Deer
Odocoileus virf^inianus dacotensis Goldman and Kellogg, 1940, Proc. Biol. Soc. Washington, 53:82, 28 June (type locality. White Earth River, Mountrail Co., North Dakota).
Occurring throughout much of North America, white-tailed deer characteristi- cally inhabit woodlands, forest edges, and thickets along streams. An esti- mated 40 million white-tailed deer in- habited the continent when European man arrived (Merwin, 1971:33). On the Great Plains, this species occupies decid- uous riparian communities, but seem- ingly avoids extensive open areas. Odo- coileus virginianus is numerous and widely distributed in the Black Hills, preferring timbered areas with adequate cover and water that support a good forb and browse understory. The main wintering ranges are in the foothills, and on the McVey Burn in Pennington County. This species is extremely adapt- able to human populations and is quite common in agricultural areas. In 1969, there were in excess of 70,000 white- tailed deer in the Black Hills proper (Richardson, 1969b: 25).
Grinnell (1875:83) wrote that this species was abundant in the vicinity of Castle Creek and the Elkhorn Prairie [Reynolds Prairie] about the head of Elk Creek, and all through the north- eastern portion of the Black Hills. He noted that 100 deer, principally of this species, were killed by the command of General Custer on 9 August 1874 cast of Custer Peak. Of 16 does killed by Dodge (1876:121) between 15 August and 10 September, only two had given birth to fawns that season. Approximately 1000 deer were taken by the Newton-Jenney Survey along Castle, Rapid, and Boxelder creeks (op. cit. :122), causing Dodge to state that O. virginianus was "more abun-
dant than any other animal" in the Black Hills. In strong contrast to these reports is that of Bailey (1888:434), who wrote that "the black-tail [mule deer] is very common through the Hills but the white- tail seems to be absent. I can find no one who has ever seen this deer here." Evi- dently, Bailey was in error in his obsera- tions, or confined his queries to the foot- hill region, because notations by W. W. Granger in 1894 agreed with the previous accounts (J. A. Allen, 1895a:263).
Odocoileus virginianus dacotensis differs from O. v. ochroura of western Wyoming (Long, 1965:714) and O. v. tuocroiirus to the east (Kellogg, 1956: 44) in average larger size, heavier den- tition, and paler color. However, there seems to be extreme variation in size within the geographic range of O. v. dac- otensis. Evidently, white-tailed deer in the Black Hills region also are quite variable and may represent a broad zone of intergradation with the adjacent sub- species. Jones (1964:319) felt that the alleged differences between subspecies did not appear to be great. The enigma remains due to a lack of specimens for examination.
Grinnell (1875:83) indicated that the differences in size between the white- tails in the Black Hills and those on the prairie were so great that hunters of the region thought the Black Hills deer to be a different species than that to the east. Dodge ( 1876:130) reported: "a few days previous to the arrival of General Cook, I killed near Rapid Creek two enormous bucks, each of which, after disembowel- ing, and having head and legs cut off, weighed nearly one hundred and thirty pounds. In the hunt on Boxelder [with General Cook], one of the guests killed a buck just as old and just as fat as those killed by me on Rapid, but which, dressed in the same way, weighed scarcely forty pounds." Dodge indicated that both the large and small forms were found together in the same herd, as well as animals representing gradations in size between the mentioned extremes. Grinnell wrote as follows to Seton
140
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
(1929b: 321): "In 1874 when with Cus- ter's expedition to the Black Hills, L. H. North and I killed in the Black Hills three or four extraordinarily small deer." Numerous authors went on record in Forest and Stream, and Recreation maga- zines during the period 1895 to 1900 as having seen small white-tailed deer in the Black Hills. Measurements of a pair of diminutive white-tail antlers from In- yan Kara Mountain, found in 1890 by Robert Ansley (Seton, 1929b: 316) are followed by those of a similarly-sized pair found in the Bear Lodge Mountains by Vernon Bailey in 1913 (USNM 202513): distance from burr to tip of antler, 8 inches, 10 inches; greatest dis- tance between any two corresponding points, 10 inches, 8.5 inches; distance from base of one antler to base of second antler, 2 inches, 1.75 inches; distance from tip of one antler to tip of other antler, 8.5 inches, 4.75 inches. A few unusually small adult white-tails still are killed by hunters each season in the Black Hills (Over and Churchill, 1945: 51; Bever, 1957:4).
Nutrition may be a possible explana- tion of the problem. Both white-tailed deer and mule deer in the Hills are smaller in body size than are prairie rep- resentatives of the same respective sub- species (Berner, 1953c: 12). Over-popu- lation and over-utilized range are char- acteristic of the Black Hills herd, but not of the prairie herd. Food habits are essentially the same for mule deer and white-tailed deer in the Black Hills, and have been reported by Hill and Harris (1943), Hill (1946), Gastler et al (1951), Bever (1954), and Schneeweis (1968). As determined by pellet counts, deer utilize mixed aspen-pine communi- ties to a greater extent than pure stands of either pine or aspen ( Kranz and Peter- sen, 1970:5). Important browse species ( based on observation, surveys of browse utilization, and analysis of stomach con- tents of 320 deer taken in the northern section of the Hills) are as follow: ground juniper, crcx^ping juniper, (juak- ing aspen, bur oak, ponderosa pine, paper
birch, hawthorn {Crataegus succuhnta), hop hornbeam, redroot, hazelnut (Corij- lus cornuta), Oregon grape, bearberry, wolfberry, snowberry (Symplwricarpos canadensis), Juneberry {Amehncheir canadensis), serviceberiy (A. spicata), chokecherry, and russet buffaloberry (SJiepJierdia canadensis). Forbs utilized by the Black Hills deer include pea vine (Lat])yriis ochroleucus) , American vetch {Vicia americana), blue aster (Aster laevis), meadow sweet, wild pea (Lu- j)inus argenteus), pussy toes (Antenna- ria microphyUa) , wintergreen (PyroJa sp.), alfalfa, red clover, wild rose, soap- weed, and various mushrooms, grasses and lichens ( Usnea sp. ) . Kentucky blue- grass comprised over 90 percent of the grasses consumed (Hill, 1946:49).
Progulski and Duerre (1964:27) found that deer in the Hills were most active between one and five hours after sunset. They indicated (op. cit. -.31) that activity was most concentrated in lush meadows that were interspersed with brushy cover along streams. Rich- ardson and DeMarce (1967:8) analyzed the numbers and distribution of deer in the Black Hills in 1966 and 1967, based on a 365-day accumulation of deer pel- lets. They found 19.7 percent of the deer occurred in the northeastern section, 47.7 percent in the northwestern-central sec- tion, 22.0 percent in the southeastern- central section, and 10.6 percent in the southeastern section of the Hills.
Mr. L. Dunn of Sturgis shot a white- tailed buck with palmated antlers near Moskee, Crook Co., Wyoming in 1965; there ^^'ere 13 points on a side and the antlers much resembled those of a small moose (Lee, 1965:19). In the autumn of 1968, Dr. Calvin Schad shot a white- tailed buck in the Black Hills that had three antlers: a normal four-point antler on the left side of the skull, and two three-point antlers, arising separately, on the right side (South Dakota Con- servation Digest, 36:13, April, 1969).
Rut begins early in November in the Black Hills, reaching its maximum dur- ing the latter part of the month. Does
TURNER: MAMMALS OF THE BLACK HILLS
141
first breed as yearlings and usually bear twin fawns from late May to early July. On one occasion I witnessed a doe with three fawns near Custer. Mule deer- white-tailed deer hybrids have been re- ported from the Hills but none have been authenticated (Bever, 1957:5).
During late summer and early au- tumn of 1952 and 1956, outbreaks of epizootic hemorrhagic disease (EHD) killed hundreds of white-tailed deer and an occasional mule deer in western South Dakota, including Meade, Pennington, Custer, and Fall River counties (Shope et al, 1960; Pirde and Lay ton, 1961; Trainer, 1964). Of the two viral strains known to be causative agents, the "Black Hills strain" is far less virulent than the "South Dakota strain" that causes epi- zootic outbreaks among prairie deer (Parikh, 1970:1). These EHD-causing strains currently are under investigation by virologists at South Dakota State Uni- versity. Deaths by automobiles is another mortality factor. For example, more than 30 deer were killed on highways in the northern Hills from 16 September to 12 October 1952 (Harris, 1952c: 16).
Specimens examined (29). — SOUTH DA- KOTA: Lawrence County: 3 mi S, 3.5 mi W Spearfish, 1. Pennington County: 16 mi W HiU City, 6600 ft, 1; 5 mi W Hill City, 2; near Pactola, 1 (USD); 6 mi NE Keystone, 2; 4 mi N, 2 mi E Keystone, 4800 ft, I; 20 mi N Elk Mountain, 1 (USNM). Custer County: Pine Creek, 1 mi N Harney Peak, 1 (SDSU); Elk Mountain, 2 (USNM).
WYOMING: Crook County: Bear Lodge Mountains, 13 (USNM); Lost Canyon, 5000 ft, 1 (UW); unspecified locality, 2 (AMNH). Weston County: vmspecified locality, 1.
Additional records.—SOUTU DAKOTA: Lawrence County: Elk Creek (Harris, 1952b: 12). Custer County: Bull Springs, 14 mi NW Custer (Lamster, 1943:14). WYOMING: Crook County: Moskee (Lee, 1965:19). Crook's Tower ( Chicago Tribune, 12 Novem- ber 1941). UNSPECIFIED LOCALITY: Black Hills (Hoffman, 1877:100).
Family ANTILOCAPRIDAE— Pronghorn
The pronghorn is not an antelope but is a member of a unique family that has evolved exclusively in North America
and is represented in the Black Hills re- gion by one Recent species. Both sexes of this species possess permanent bony outgrowths from the skull that are cov- ered with highly keratinized horny sheaths; these sheaths are shed and re- placed annually. Pronghorn are grazers and inhabit the open flat country and gently rolling upkmds of the Great Plains.
Antilocapra americana americana (Ord)
Pronghorn
Antilope americana Ord, 1815, in Guthrie, A new geograpliical, historical, and commer- cial grammar . . ., ed. 2, 2:292, 308 (type locality, unknown; noted in the original description as found "on the plains and high-lands of the Missouri").
Antilocapra americana Ord, 1818, Jour. Phys. Chim. Hist. Nat. et Arts, 87:149.
Pronghorns now are reduced in num- bers over much of their former range. An estimated 30 to 40 million once roamed over the western half of the United States (Long, 1965:716). Agri- cultural pursuits of man and settlement of former pronghorn range contributed to a decline in the abundance of this species (Nelson, 1925:3). Hoffman (1877:100) indicated that a fatal epi- demic raged among pronghorns in the summer of 1873, and that J. A. Allen estimated that from three-fourths to nine- tenths of the population may have been destroyed in some areas. Coincident with this epidemic, an epizootic disease killed most of the government stock and Indian ponies in western South Dakota. Hoffman {loc. cit.) noted that "If the horse epidemic was not the cause of fatality among the antelope, it is at least in very remarkable coincidence." Nearly 64 percent of the Indians at this time were affected with cerebro-spinal menin- gitis, and suffered a 10 to 12 percent mortality. The Indians procured their drinking water from the same pools where their ponies drank.
On 16 October 1804, Indians told Meriwether Lewis and William Clark that the pronghorn were then on their
142
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
fall migration west to the "Black Moun- tains" to spend the winter ( Bailey, 1927 : 28). Hayden (1862:150) wrote: "In the beginning of winter they [pronghorn] may be seen for days following each other in files (if not disturbed) on their way towards the Northwest, leaving the prairie for the more rugged portions of the country near the Black Hills or the foot of the mountains. In the spring, usually about March, they may be seen returning again, and distributing them- selves over the open prairie." Seton (1929b: 421) also reported that prong- horn of the open country flock to the Black Hills "from all points on the com- pass" to winter.
In his review of the status of the pronghorn. Nelson (1925:3) recorded 11 bands, totaling an estimated 680 indi- viduals, as being resident in South Da- kota, and Berner (1954:2-3, 1955:2-3) summarized the history of the prong- horn in South Dakota. At present, spring inventory of pronghorn is carried out by the South Dakota Department of Game, Fish and Parks each year. The aerial count, conducted from 3 June to 8 July 1970, indicated that the population in western South Dakota numbers about 25,000 individuals, representing a six per- cent increase over the 1969 estimate (West, 1971:1). A census of pronghorn in counties east of the Missouri has not been attempted for several years, but 302 individuals were killed by hunters in these areas in 1968 (West, 1970:2).
Present rangeland management in western South Dakota is oriented toward grasses and other graminoid plants. Be- cause native woody range plants and forbs are high in fats, oils, carotenes, protein, and other constituents necessary for winter maintenance of both big game and domestic livestock. West (1971:4) suggested the need for re-establishment of these browse-plants species. The cur- rent population of pronghorn on that portion of the prairie adjacent to the Black Hills is about 82 percent of esti- mated carrying capacity ( see Table 28 ) .
In managing pronghorn, the Depart- ment of Game, Fish and Parks divides sections of South Dakota into "Hunting Units" that can be censused and adminis- tered independently. Units that surround the Black Hills are as follow: Unit 3, western Butte County (just north of the Hills); Unit 4, eastern Butte and north- western Meade counties; Unit 9, southern Meade, central Pennington, and western Custer counties; Unit 14, eastern Fall River County; Unit 6, western Fall River and southwestern Custer counties. Table 28 summarizes the composition of the 1970 pronghorn populations in these re- spective units (see West, 1971: Tables land 2).
A herd of about 300 pronghorn cur- rently is maintained in Wind Cave Na- tional Park. Dr. W. T. Hornaday of the Bronx Zoo, and the Boone and Crockett Club, contributed the monev for the initial restocking of the Park in 1914.
Table 28. — Estimated composition of the 1970 populations of Antilocapia americana in Hunting Units that border the Black Hills (West, 1971 ).
Hunting Unit
|
•« to |
<0 in |
Si |
n2, ij O |
|
|
umb buc |
umb kids |
o XI |
||
|
Z^ |
Z o |
Z O |
Q |
o
•43
O N (In -S
be
;>>
>. O
u S
3 480 950 970 100:102 2400 3500
4 533 886 806 100:91 2225 3350
6 315 516 489 100:95 1320 1600
9 288 432 480 100:111 1200 1500
14 275 436 389 100:89 1100 1000
Total or average .. 1891 3220 3134 100:97 8245 10950
TURNER: MAMMALS OF THE BLACK HILLS
143
With permission of the Canadian authori- ties, 13 six-month-old kids were pur- chased from Mr. Blazier of Brooks, Al- berta, for $125.00 eaeh. These were reared for a short time, then were liber- ated in the Park that October (Nelson, 1925:9; Grinncll, 1929:139; Wind Cave National Park files). Disease and preda- tors reduced the number to eight by 1915, and nine more were purchased from Alberta in 1916. Coyotes and bob- cats continued to take a high toll of the pronghorn until predator)' animal hunt- ers were detached to the Park by the U. S. Biological Survey; these hunters eliminated sexeral hundred predators (Nelson, 1925:17). By 1924, only six animals ( all does ) survived, but a young buck was captured near Bisner, Nevada, and released in the Park in July. The herd numbered only 28 by 1929 (Grin- ncll, 1929:140), but 40 individuals sur- vived the disease and drought of the 1930's. Pronghorn finally were well- established in the Park by 1948 and num- bered about 150 animals, in spite of the loss of 35 individuals to poachers in the previous year (Cahalane, 1948:256). It is probable that control of carnivores in the region no longer is of merit owing to the natural balance that now exists between predator and prey.
Management programs have involved transporting live-trapped individuals to areas of low population density for re- lease. These introductions have been numerous and difficult to trace. Twelve kids were introduced into Custer State Park from Indian Creek Flats, Butte County, in 1916, and 10 more were added in 1919 ( South Dakota Conservation Di- gest, 36:3, 1969). Eight pronghorns trapped in Meade County were released in Custer State Park on 4 February 1952 (Berner, 1952:8). Fift\'-one pronghorn were transplanted from Custer State Park to Leola Hills, near Milbank, Grant County, in 1959 (South Dakota Conser- vation Highlights, 1960), and 126 were transferred from Butte Count}^ to Day and Edwards counties in 1961 (South Dakota Conservation Highlights, 1962).
Numerous pronghorn were captured in northwestern South Dakota for release on the Lower Brule Indian Reservation, Lyman County, and on the Igloo Black Hills Ordnance Depot, Fall River County, in 1963 (South Dakota (Conser- vation Highlights, 1964). One hundred and twenty pronghorn were live-trapped in Wind Cave National Park in 1963 for release on the Lower Brule Indian Res- ervation, on private lands in Gregory County, and for the Kansas Game and Fish Commission for release in southern Kansas ( Wind Cave National Park files ) . These are but a few instances of artificial mixing of the herds in South Dakota.
On 2 September 1964, the "Head- quarters Fire" burned approximately 4000 acres of Wind Cave National Park. The kid crop of the next year was much reduced from that of previous years. Pronghorn moved into the burned area and fed mainly upon pricklypear cactus just prior to the peak of the reproductive season. False gromwell (Litlwspermum multiflomm) is known to inhibit ovula- tion in some grazing mammals, and Ranger-Naturahsts at the Park speculate on the possibility that cacti, from which the spines have been burned, might have had a similar eff^ect on the reproductive potential of pronghorn ( from various re- ports on file at Wind Cave National Park). Breeding takes place in Septem- ber and young, usually twins, are born in late May and June.
Specimens examined (10). — WYOMING: Crook County: Sundance, 7 (USNM). Weston County: 3 mi S, 1 mi E Newcastle, 1; imspeci- fied locality, 2.
Family BOVIDAE — Bison, Sheep, AND Goats
This family of large herbivores is represented in the Black Hills by thi-ee species. Two of these formerly were na- tive to the region and have been reintro- duced, one being replaced by a non- native but similar subspecies. The third representative is a non-indigenous spe- cies established by man.
144
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Bison bison bison (Linnaeus) Bison
Bos bison Linnaeus, 1758, Systema naturae, ed.
10, 1:72 (type locality, "Quivira," central
Kansas — see Hershkovitz. Proc. Biol. Soc.
Washington, 70:32, 28 June 1957). B[ison'\. bison — Jordan, 1888, Manual of the
vertebrate animals . . ., ed. 5, p. 337.
The number of bison that once roamed the plains of North America has been estimated between 60 and 100 mil- Hon by various authors. The bison evi- dently was common on the broad, un- dulating prairies surrounding the Black Hills in the early 1800's, but at the time of the first exploration of the region by European man, it was rare to see even an occasional wanderer. Bison had wholly disappeared east of the Missouri Biver prior to 1870, but great herds roamed the Coteau des Prairies west of the James River in South Dakota as late as 1866 (J. A. Allen, 1877:538). This species was decimated in the region be- tween the Grand and Cheyenne rivers about 1869, although occasional strag- glers frequented the plains toward the Black Hills until somewhat later (Hoff- man, 1877:101).
When Lt. G. K. Warren (1859:630) and his command traversed the western edge of the Black Hills in September 1857, they encountered a large force of Hunkpapa and Miniconjou Sioux in the vicinity of Inyan Kara Mountain, Crook County, Wyoming. Warren gave the following account: "In the first place, they [the Indians] were encamped near large herds of buffalo, whose hair not being sufficiently grown to make robes, the Indians were, it may be said, actually herding the animals. The intention of the Indians was to retain the buffalo in their neighborhood till their skins would answer for robes, then to kill the animals by surrounding one band at a time and completely destroying each member of it." Fearing that Warren's expedition would disperse the bison, the Indians forced it to turn back southeastward. Most writers have attributed the rapid extermination of the bison over south-
western South Dakota and adjoining portions of Wyoming to white man, rather than to the Sioux Indians. If the Sioux often hunted in the manner de- scribed by Warren, then their contribu- tion toward the decimation of bison in the Black Hills region was great indeed.
General W. F. Raynolds (1865:27), in the course of leading an expedition to explore the Yellowstone in 1859, caught first sight of the "lords of the prairie" as his command approached the Black Hills. When military posts just to the west of the Black Hills were abandoned in 1868, bison were still abundant, 50 tons being killed for garrison use in one day; yet, by 1871 no bison survived in eastern Wyoming (J. A. Allen, 1877:544). Grinnell ( 1875 : 84 ) , while accompanying General Custer in 1874, observed no live bison in the Black Hills, but found a few skulls with the hide still attached, indi- cating recent presence of this bovid. Much later, Fryxell (1926:103) also dis- covered bison remains in the Black Hills.
Bison in the Black Hills today are the result of reintroduction. In 1913, the National Bison Society provided $26,000 with which to purchase bison to restock the Wind Cave National Game Reserve. W. T. Hornaday secured the nucleus herd of 14 animals (seven bulls and seven cows ) from the Bronx Zoo through the New York Zoological Society on 28 November 1913. These bison were orig- inally acquired by the zoo from WilHam C. Whitney of the October Mountain Game Preserve in the Berkshire Hills near Lenox, Massachusetts, in 1903. Whitney, in turn, purchased them from H. K. Gliddon, of Moosehcad Ranch at Jackson, Wyoming; one of the bulls had been captured wild in the panhandle of Texas by Colonel Charles J. "Buffalo" Jones. Two bulls and four cows from Yellowstone National Park were added to the herd in 1916.
The Custer State Park bison herd, now the second largest in the United States, was started in 1914. The South Dakota State Legislature appropriated $15,000 to stock the reserve with 36 ani-
TURNER: MAMMALS OF THE BLACK HILLS
145
mals (Six bulls, 18 cows, and 12 calves) that were purchased from the James 'Scotty" Philip estate, Fort Pierre, South 'Dakota. This herd originated from five cahes captured along the Grand River near Fort Bennet, in northwestern South Dakota, by Peter Dupree in 1882. The Dupree herd increased to 60 by 1900, when Philip purchased the entire stock. An additional 60 bison were obtained by the State Park from the Pine Ridge In- dian Reser\ation, Rosebud, South Da- ikota, in 1951 (South Dakota Conserva- tion Digest, 20:23, December, 1953). I Presently, approximately 350 animals 'range over 27,220 acres in Wind Cave National Park, and about 1550 animals range over 72,000 acres in Custer State Park. These numbers are maintained by culling and selective harvesting. For ex- ample, 276 bison were auctioned by Cus- ter State Park in 1968 at a live sale and 138 others were butchered (South Da- kota Conservation Digest, 36:29, Decem- ber 1969). Each autumn, usually in October, the bison in Wind Cave Na- tional Park are rounded up by use of helicopters, horses, and vehicles, herded into a corral, branded, and treated for Bangs disease (Brucellosis). Excess ani- mals are removed for slaughter and sale in cooperation with the State of South Dakota ( South Dakota Conservation Di- gest, 20:3-5, 7; Griffith, 1958:16-19). In 1968, 10 hunting permits ($500 each) were issued to hunt bison in Custer State Park.
Throughout most of the year, bulls remain solitary, or in small groups. About late July, the bulls begin to join the cow-calf herd and mating occurs, usu- ally in August. Calves are born between mid-April and June, after a gestation period of about nine months. Presently, the Wind Cave herd is being studied extensively by Milo Schulte, Iowa State University, Ames, Iowa. Garretson (1938) and Roe (1951) have written valuable accounts concerning the past history, disti'ibution, and natural history of l)ison.
Specimens examined (2). — SOUTH DA-
KOTA: Custer County: "Wind Cave Game Reserve," 1 (USNM). Vail River County: Hot Springs, 1 (USNM).
Additional records (see text also). — SOUTH DAKOTA: Custer County: French Creek (Grinncll, 1875:84).
Oreamnos americanus missoulae
J. A. Allen
Mountain Goat
Oreamnos montanus m,issoidae J. A. Allen, 1904, Bull. Amer. Mus. Nat. Hist", 20:20, 10 Feb- ruary (type locality, Missoula, Missoula Co., Montana).
Oreaninos americanus missoulae — Hollister, 1912, Proc. Biol. Soc. Washington, 25:186, 24 December.
Mountain goats are not native to the Black Hills (Swift, 1941:441) and other- wise have not become established east of the Continental Divide. In February 1924, U. S. Senator Peter Norbeck and State Game Warden L. C. Hawley ob- tained six goats, four females ( a yearling and three adults) and two males (a yearling and an adult), from near Banff, Alberta, Canada (Gilhland, 1968a :3). These were placed in a 20-acre pen in Custer State Park, but an adult female and a yearling male escaped the first night. Warden Hawley indicated to Hanson (1950:21) that no young were born to the captive goats, but Harmon ( 1944 : 149 ) stated that the remaining four increased to eight by 1929, at which time all escaped (when a tree fell across the pen fence) to the rugged Elkhorn country near Harney Peak. The herd increased to 300 by 1949, and the popu- lation has remained at about that level to the present time. Goats were trans- planted to Spearfish Canyon, north of Savoy, several years ago, but none has been observed there since then (South Dakota Conservation Digest, 22:11, July, 1955).
Oreamnos americanus is stenoecious in the Black Hills, inhabiting only the Mount Rushmore-Needles-Harney Peak area in the southern part of the crystal- line central basin, where moss- and lichen-co\ ered granitic ridges are thrust upward nearly perpendicular to the sur-
146
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
rounding forest. The range of this spe- cies in the Hills comprises approximately 32,000 acres, of which only 2080 acres are "primary range" (Schara, 1967:22). Population density on the primary range is about 12 goats per square mile, and approximates four goats per square mile on less desirable range (Hanson, 1950: 27).
Harmon (1944:149-151) and Hanson ( 1950:55-63) reported on the food habits and natural history of the Black Hills mountain goat herd. Feeding takes place at short, irregular intervals throughout the day, but most intensive foraging oc- curs from late afternoon to dark. Food plants generally include mosses, lichens, grasses, bracken fern {Pteridhwi aqtiili- mim), yarrow (Aclnllea lamilosa) cur- rant (Rihes inehrians), fleabane {Erig- eron sp.), bearberry, serviceberry, choke- berry, hazelnut, buffaloberry, meadow sweet, paper birch, willows, quaking aspen, ponderosa pine, white spruce, and ground juniper. The most important food of all seasons, both in preference and in quantity consumed, is the tree lichen known as "old-man's beard" ( Usnea harbata). Feeding sites and food habits vary with season. In spring, the goats feed on succulent graminoid vege- tation in small grassy parks, near rocky areas, along the bottoms of canyons. In summer and autumn, they forage among small coves of aspen and birch scattered over slopes and ridges. In winter, the goats retire to the timber, or descend to protected areas such as the gulches of Grizzly and Pine creeks, there competing with deer and elk for available food. Winter foods consist of 60 percent mosses and lichens, 20 percent bearberry, 10 per- cent pine twigs and needles, and 10 per- cent miscellaneous browse, grasses, and weeds. Goats frequently visit salt blocks placed atop Harney Peak by the U. S. Forest Service.
In the Black Hills, district seasonal migrations are limited or nonexistent, aside from short altitudinal movements correlated with procurement of food (Bever, 1955:4). Mature goats in the
Hills evidently establish individual home ranges and remain within these limits the year around (Hanson, 1950:74). Al- though home ranges overlap one another to varying degrees, the goats seldom gather into groups except during rut.
Mating usually occurs in October and November. After a gestation period of about 180 days, one (usually) or two (occasionally) young are born in April or May in a secluded place amid dense timber that is intermingled with rock- piles (Hanson, 1950:42). Kids are weaned the following August, but may accompany the mother until the next young is born. Reproduction does not occur until two years of age. Of 233 goats sampled from Julv 1948 to February 1949, Hanson (1950:28) reported 138 adults, 36 yearlings, and 59 young.
Shed of the long outercoat begins in mid- April; the worn outer fur is com- pletely shed by early July. The shorter undercoat molts in the latter part of this period. A new outer coat is fully grown by mid-September (Hanson, 1950:36- 38).
A recent survey by Arthur H. Rich- ardson (pers. com.). State Game Biolo- gist, indicated that the browse plants have been severely overused, with pre- ferred food species being drastically re- duced in the primary range of the goats. Trapping operations indicated that the general health of the herd was declining rapidly. In order to combat a feared population crash, the first mountain goat season was held in 1967. Forty-six goats were taken by hunters in the next two years, several of which were large enough to be registered with the Boone and Crockett Club. On 8 March 1956, a large male scoring 53K points, was found dead along Grizzly Creek, 3 mi E Harney Peak (South Dakota Conserva- tion Digest, 24:5, August, 1956).
Visceral samples colected from 28 goats in the 1967 and 1968 hunting sea- sons revealed that the population was heavily infested with endoparasites (South Dakota Conser\ation Digest, 36:5, December, 1969). Three kinds of
TURNER: MAMMALS OF THE BLACK HILLS
147
cestodes {Moniezia J)cneden\, Taenia hijdatigena, and Thi/sanosoma aciinoides in the intestines) and 11 kinds of nema- todes (Pwtostronyhis nishi and P. stUesi in the lungs; 0.stcrtaij,ia circuincincta, O. ostertagL O. trifitrcata, TricJiostrogyhis axei, and T. coliuiibriformis in the abom- asa; Neniatodinis Jwlvetianus and N. macidosus in the small intestine; Oesoph- agastonium vemdosum and TricJiuris sp. in the caeca) ha\'e been identified from these visceral samples (Boddicker and Hugghins, 1969; Boddicker, 1970; Block, 1970- Boddicker et al, 1971). Ticks (Dermacentor andersoni) and chewing lice (Bovicola. oreatnnidis) parasitized mountain goats in the Black Hills ex- ternally (Boddicker et al, 1971:95), and residues of 0.59 parts per million of chlorinated hydrocarbons ( heptachlor epoxide, dieldrin, DDD, and DDE ) were isolated from kidney fat of 13 goats (op. d^:101).
Overcrowding of available range and heaxy lungworm infestations that result in diseases of the lungworm-pneumonia complex probably are major factors in controlling population levels of mountain goats in the Black Hills. A treatise on this species and its management cur- rently is in press (A. H. Richardson, The Rocky Mountain Goat in the Black Hills. South Dakota Dept. Game, Fish and Parks, Bull. 2, Pierre).
Specimens examined (2). — SOUTH DA- KOTA: Cttster County: Harney Peak, 1 (SDSU). Unspecified County: Black Hills, 1 (SDSU).
Ovis canadensis auduboni Merriam
Mountain Sheep
Ovis canadensis auduhoni Merriam, 1901, Proc. Biol. Soc. Washington, 14:31, 5 April (type locality, Upper Missouri, probalaly Badlands betxveen the Cheyenne and White rivers. South Dakota ) .
Mountain sheep formerly inhabited foothills and prairie breaks, as well as mountainous areas, in the western United States, but now occupy only rough buttes and canyons in the more rugged moun- tains. Ovis canadensis auduhoni, a sub-
species that now is extinct (Cowan, 1940: 542), was native to the Black Hills. In- troduction (Jackson, 1944:28) of the morphologically similar \\{\st(>rn subspe- cies, O. c. canadensis, accounts for the present occurrence of moimtain sheep in the Hills.
Honess and Frost (1942:4) indicated that O. c. auduhoni may be indistinguish- able from O. c. canadensis. Lacking specimens, I follow Cowan (1940:542- 543), who critically examined material from throughout the range of the species, in applying the name auduhoni to the native mountain sheep of the Black Hills and adjacent areas. However, Cowan (1940:543) noted that "O. c. auduhoni based as it is on slight cranial characters presented by a small number of speci- mens is to be regarded as a weak race."
In 1833, Maximillian reported that the Manitarii Indians usually went to the Black Hills and associated mountainous tracts to hunt mountain sheep, killing a hundred or more in a season (Thwaites, 1906:246). However, the maps of Maxi- millian labeled the Killdeer Mountains and the Badlands of the Little Missouri River (both in North Dakota) as the "Black Hills" (Bailey, 1927:25). Mem- bers of the Newton-Jenney U. S. Geologi- cal Survey shot a mountain sheep along a small tributary of Boxelder Creek on 20 July 1875 (Dodge, 1976:129). Seton (1929b: 535) reported that mountain sheep were essentially gone from the Black Hills by 1887, although a few lin- gered on until 1899 when the last indi- vidual about which he had information was killed. W. S. Phillips, however, in- dicated that mountain sheep ranged in the summer of 1890 from Sundance, Wy- oming, to the Inyan Kara and Bear Lodge mountains, all along the western slopes of the Black Hills (Grinnell, 1904: 3.34-335), and J. A. Allen ( 1895a: 263) re- ported a small herd in the vicinity of Harney Peak near the turn of the century. No sheep were seen by Seton ( loc. cit. ) when he xisited the Hills in the summer of 1902, but a band of about 200 O. canadensis still survived to the east in
148
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
the White River Badlands, near Porcu- pine Hill in 1909 (Charles, 1964:23). The idea that native sheep were initially prairie dwellers (Grinnell, 1928:1) and were driven to the mountains seems to be more closely associated with audu- honi than with any of the other races (Buechner, 1960:21).
A herd of eight Ovis canadensis, ob- tained from Alberta, Canada, in 1922 by U. S. Senator Peter Norbeck, was con- fined in Custer State Park, but a sore- mouth virus all but eliminated them bv 1961. A new herd was brought in from Montana about 1965 and now consists of from 50 to 70 sheep (A. H. Richardson, pers. com.). Some indi\'iduals, mainly rams, escaped into the surrounding Hills. Possibly, additional individuals from other herds introduced into nearby areas in western South Dakota may have reached the Black Hills. Twelve moun- tain sheep obtained from Alberta were released on the Claude Olson Ranch, Slim Buttes, Harding County, in 1961
(Nachtigal, 1961:4), and 22 from Colo- rado were received at the Badlands Na- tional iMonument in 1963 (Charles, 1961: 5; Conservation Highlights, South Da- kota Dcpt. Fish, Game and Parks, 1964). In January 1967, a single ewe with one lamb was observed in the Red Valley in Wind Cave National Park. A skeleton was found in the same area in the sum- mer of 1968, creating some doubt as to the survival of the ewe and lamb. On 15 January 1969, nine mountain sheep ( one adult male, seven adult females, one ju- venile male) were transplanted from Du- bois, Wyoming, to the northwest side of the In\'an Kara Mountain in sec. 24, T. 49 N, R. 63 W, Crook County, Wyoming. Eight of these ascended the mountain slope, but one female escaped across the prairie. As of 8 October 1969, the mature ram and one female died; howe\'er, four new lambs have been observed (F. C. Windsor, pers. com.).
Specimens examined (1). — SOUTH DA- KOTA: Custer Cotmtij: Wind Cave National Park, 1 (WCNP).
SPECIES OF UNVERIFIED OCCURRENCE
SPECIES INCORRECTLY REPORTED FROM THE BLACK HILLS
Sorex vagrans monticola Merriam, 1890.— Baker and Findley (1953:382) reported a specimen of the vagrant shrew from 2.5 mi N Fairburn, Custer Co., South Dakota. They wrote as follows: "We have tentatively referred this speci- men to S. t). monticola Merriam on geo- graphic grounds, although certain cranial measurements taken show the shrew to be much smaller than other shrews of this subspecies. . . ." Subsequently these same authors (Findley and Baker, 1956: 543) assigned the same specimen to Sorex nanus (see account of that spe- cies). Findley (1955:26) suggests that a segment of the ancestral S. vagrans stock might have persisted in the Black Hills during the Sangamonian interval.
Sorex vagrans obscurus Merriam, 1891.— Long (1965:522) reported a
specimen of the \'agrant shrew in the Museum of Natural History, The Uni- versity of Kansas, from a localitv 3 mi NW Sundance, 5900 ft, Crook Co., Wy- oming. There is no evidence that such a specimen e\'er existed at the Museum; however, there is an additional speci- men of Sorex cinereiis Jiaydeni beyond the number indicated by Long for the specified localit)^ Findley ( 1955 ) revised the species Sorex vagrans and listed no specimens from Crook County; yet. Long (1965:520) wrote, ". . . my findings agree with his [Findley] concerning S. vagrans in Wyoming." Thus it appears that Long's report of the vagrant shrew occurring in the Black Hills is in error.
Myotis evotis evotis (H. Allen, 1864.) — The long-cared m\'otis has been re- ported from the Badkmds to the south- east of the Black Hills and from Hard- ing County to the north. It also is known
TURNER: MAMMALS ()i< HIE BLACK HILLS
149
from much of Wyoming, excluding the northeastern portion of the state. Pre- vious reports of M. evotis from the Hills were erroneous. Of three specimens listed by Stebler (1939:389) from Custer State Park, South Dakota, one is Myotis tlujsanodes, and the other two are M. keenii. One reported from 1.5 mi E Buckhorn, Weston Co., Wyoming ( Long, 1965:532) also is M. tJujsanodes.
Geomys bursarius lutescens Merriam, 1890.— Long (1965:613) reported a specimen of the plains pocket gopher from 0.5 mi N and 1 mi E Beulah, 3550 ft. Crook Co., Wyoming, that allegedly w as housed in The University of Kansas, Museum of Natural History. I have ex- amined the museum catalogue and spe- cies card files, the field notes and cata- logues of members of the field party that collected at this site, and all specimens of Geomys hursarius from Wyoming in the Museum's collections. Nowhere is there evidence of the indicated specimen and Long's report thus may be consid- ered as being erroneous. Representatives of this species have been taken on the flood plains of the Cheyenne River, on the southern periphery of the Hills. There are three specimens from 1 mi E Edgemont and two specimens from 6 mi S Hot Springs (UMMZ), Fall River Co., South Dakota. D. G. Adolphson recently retrieved several skulls of this species from pellets of a barn owl nesting under the Hat Creek bridge on Highway 71, 15 mi S Hot Springs, in Fall River County (Martin, 1971a, b).
Phenacomys intermedius intermedius
Merriam, 1889.— Cones and Allen (1877: 216-217) recorded a specimen of Arvic- ola (Pedomys) haydeni (USNM 3056) taken from the "Black Hills." They con- sidered the individual as being peculiar and accredited it to being "a young of the year." Upon reexamination, J. A. Allen (1894a:331) assigned the specimen to a new species of Phenacomys (P. fruei, which is in the synonymy of P. interme- dius). In actuality, the indixidual was collected by W. A. Hammond on 10 Au-
gust 1857 "along a wagon road to Bridger Pass" in the region now known as the Laramie Mountains, Wyoming, and the heath(>r \'ole does not occur in the Black Hills.
Zapus princeps princeps J. A. Allen, 1893. — When composing the general list of mammals collected by W. W. Granger in 1894, J. A. Allen (1895a:262) included 7Mpus princeps (actually, these were Z. Jmdsoniiis) among those mammals taken in the Black Hills, but not from the Badlands. Yet, in the species ac- counts {op. cit. -.266), the only western jumping mice obtained by Granger were listed as from Corral Draw in the Bad- lands. I know of no representatives of this species from western South Dakota or northeastern Wyoming, where only Z. hudsonius occurs.
Spermophilus lateralis lateralis (Say,
1823). — The golden-mantled ground squirrel does not inhabit the Black Hills at the present time. Coues and Allen (1877:818) listed a male of this species that was collected in the "Black Hills" on 21 July of 1859 by W. S. Wood, who accompanied the expedition of Lieuten- ant F. T. Bryan through the Laramie Mountains, and who, to my knowledge, did not enter the Black Hills proper. Spermoplidtis lateralis does occur about 110 miles to the southeast, in northeast- ern Albany County, Wyoming (Long, 1965:588).
SPECIES OF UNCERTAIN STATUS IN THE BLACK HILLS REGION
Didelphis marsupialis virginiana Kerr, 1792. — A young opossum that was cap- tured in northern Converse County, Wy- oming, by Mr. Hans W. Larsen, subse- quently escaped from a cage in New- castle, Weston County, in 1962 (Long, 1965:515). Individuals of this species have been observed in recent years in Mellette and Todd counties. South Da- kota, and a specimen was captured in 1958 at Alliance, Box Butte Co., Ne- braska (Jones, 1964:58). The opossum
150
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
now may be expanding its range into the Black Hills from the east by way of the deciduous riparian communities of river systems.
Sorex nanus Merriam, 1895. — The dwarf shrew has been reported just east of the Black Hills on the basis of a par- tially decomposed specimen recovered from a stubblefield 2.5 mi N Fairburn, Custer Co., South Dakota. The individ- ual first was reported as S. vagrans (Baker and Findley, 1953:382), but later was assigned to S. nanus (Findley and Baker, 1956:543). The partial skull was reexamined recently by Robert S. Hoff- mann and me, and we concur with the assignment of this specimen to S. nanus. On 3 March 1970, Don Polen, South Dakota School of Mines and Technol- ogy, captured a female of this species, along with two female Sorex cinereus, in a grassy field about 10 miles east of the Hills, at a place 3 mi E Boxelder, Pennington Co., South Dakota (Martin, 1971b). External measurements of this specimen were as follow: total body length, 82; tail length, 35.5; and hind foot length, 9. Martin (1971a, b) also re- ported a left mandible of S. nanus from a barn owl pellet picked up 15 mi S Hot Springs, Fall River County. Dwarf shrews usually are found in subalpine and alpine rockslides, at elevations of 8000-10,000 feet; thus, the three speci- mens from South Dakota are unusual both in the low altitude at which they were taken and in the habitat thev occu- pied (Hoffmann and Taber, 1960:2.33).
Ecological distribution of the dwarf shrew seems to be independent of the availability of ground water. These ani- mals seldom have been taken in snap- traps, but recent use of pitfalls has proven to be a successful method of ob- taining specimens (Brown, 1967:622). Unfortunately, the majority of trapping in the Hills region has been by use of snap-traps and dry rocky areas have not been investigated thoroughly.
The ancestor of S. nanus "may have occurred in the Black Hills and isolated mountains of Arizona and New Mexico
during the Sangamonian interval and remained in these general areas during the Wisconsin age" (Findley, 1955:28). Montane habitat in the Black Hills pre- sumably would be suitable for the dwarf shrew and future application of pitfall sampling techniques should produce specimens from this region.
Myotis grisescens A. H. Howell, 1909. —On 21 August 1968, Mr. Tim Joseph of Menden, Missouri, captured a banded (B6-02491) male bat (presumably a M. grisescens) in a small cave near Savoy, Lawrence Co., South Dakota (Gunier, 1971:5). The bat previously had been captured near Tipton, Moniteau Co., Missouri, in 1967. It was transported to Higginsville, Lafayette Co., Missouri, about 65 miles northwest of the site of capture, to a place where it was released in a region where there are no known caves ( Gunier, pers. com. ) .
The intervening distance between Higginsville and Savoy is approximately 640 linear miles across the semiarid Northern Great Plains. Presumably, this environ would be a substantial barrier to a ca\c-inhabiting species such as M. grisescens. At no previous time has this chiropteran been taken anywhere near South Dakota. The gray bat definitely is not a normal component of the Black Hills mammalian fauna and identifica- tion of bat B6-02491 and the circum- stances of its recapture thus remain sus- pect. At best, the record represents ab- normal wandering of a displaced mam- mal that was liberated in strange sur- roundings.
Euderma maculatum (J. A. Allen, 1891). — The rare spotted bat has been recorded from south-central Montana and from Big Horn County in northern Wyoming. This species possibly will be found along the deep moist canyons that are bordered by exposed limestone cliffs in the northern portion of the Black Hills.
Lepus americanus seclusus Baker and Hankins, 1950. — The snowshoe hare evi- dentlv does not occur in the Black Hills,
TURNER: MAMMALS OF THE BLACK HILLS
151
even though the distribution maps of Burt and Grossenheidcr (1964:212) and notations of Seton (1929c:739) indicate its presence there. Forest Rangers sta- tioned at Newcastle have "heard of snowshoe rabbits" around Moskee, Crook Co., Wyoming, but I know of no speci- mens from the Hills. Baker and Hankins (1950:64) stated: "Immediately east- ward of the Bighorns, there are no snow- shoe rabbits; at least none has been reported from any of the higher areas where they might be expected to occur, as for example, the Black Hills." Mon- tane habitats of the Hills presumably would be suitable for this species, how- ever.
Lepus calif ornicus melanotis M earns, 1890.— Black-tailed jackrabbits rarely may occur on the fringes of the Black Hills. Game Warden H. J. Brockley and Forest Ranger Duke Kocer both have indicated an abundance of blacktails on the flats to the east and south of the Hills. I found one killed on a road a few miles southeast of Hot Springs, Fall River County, in June 1968. Other speci- mens ha\e been reported nearby as fol- low: 1 mi N and 3 mi E Orin, Con\'erse Co., Wyoming (Long, 1965:552), and 25 mi E and 20 mi S Rapid Citv, Custer Co., South Dakota (Findley, 1956:2). In 1956-57, fur dealers from western and southwestern South Dakota reported skins of 8500 black-tailed jackrabbits (South Dakota Conservation Digest, 24: 67, June 1957). The majority of these probably were from Nebraska and Wy- oming, but there is evidence of increased numbers near the Hills.
Spermophilus spilosoma obsoletus
Kennicott, 1863. — When Kennicott (1863:157-158) described Spermophilus obsoletus, he did not designate a holo- type, but had at hand seven cotypes on which he based the description and name (Lyon and Osgood, 1909:168-169); A. H. Howell (1938-130) later selected one of these as the lectotype. Among the cotypes is a specimen, received by the Smithsonian Institution from Lieu-
tenant G. K. Warren, that was collected by F. V. Hayden in the Black Hills of Nebraska Territory [now South Dakota] sometime in the period 1857-58. This specimen now is housed in the Museum of Comparative Zoology, Harvard Uni- \ersity (MCZ 4817, formerly USNM 3252/479.5— G. M. Allen, 1931:252), al- though I ha\'e not examined it.
Ground squirrels of this species pre- fer sandy soils having sparse or low- growing vegetation. A single specimen of S. s. ohsoJetns has been taken along the south fork of White River in south- western South Dakota (A. H. Howell, 1938:131) and 12 others have been re- ported from the three southeasternmost counties in Wyoming, viz. Platte, Goshen, and Laramie (Long, 1965:577). Despite extensive collecting, no addi- tional specimens have been obtained in the Black Hills. Several Ranger-Natural- ists at Wind Cave National Park re- ported seeing Spermophilus spilosoma in the upland prairie regions of the Park in the summer of 1968. Although I sus- pect that their identifications were cor- rect, I did not observe or take any speci- mens of this species while conducting field work within the Park.
Perognathus flavus piperi Goldman, 1917. — The silky pocket mouse has been taken in northwestern Nebraska, and at the type locality 23 mi SW Newcastle, Weston Co., Wyoming (Long, 1965: 616). Cones (1875:303) and Cones and Allen (1877:518) reported a specimen of "CricetoclifAis favus' (USNM 3097) from the Black Hills. I have been unable to locate this indi\ idual in order to verify the identity, but know of no other speci- mens of this species from the region. However, the fine sandy loam soils along the South Dakota-Wyoming border pre- sumably would provide a suitable en- vironment for P. flavus.
Reithrodontomys montanus albescens
Gary, 1903. — The plains harvest mouse occurs exclusi\ely in upland habitats, preferably on sandy soils. The mixed- grass prairie uplands of Wind Cave
152
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
National Park and some of the drier draws of the foothills would seem suit- able for this species. Although no speci- mens have been obtained from the Black Hills proper, R. montcinus has been taken from localities near the Hills in Harding, Meade, Lawrence, and Fall Ri\er coun- ties. South Dakota, and in Campbell and Niobrara counties, Wyoming. For example, on 4 September 1968 I cap- tured a female plains harvest mouse along the grassy roadside of Highway 79, 1 mi N and 5.5 mi E Hot Springs, im- mediately adjacent to the Black Hills.
Onychomys leucogaster arcticeps
Hollister, 1914. — A male northern grass- hopper mouse was obtained at New- castle, Weston Co., Wyoming on 20 May 1894 by Vernon Bailey. Newcastle is immediately adjacent to the Black Hills and supports elements of both the Hills fauna and the plains fauna. Additional specimens have been taken in South Da- kota on the peripheiy of the Hills at Buffalo Gap, Custer County; Spring Creek, Pennington County (MHM); 5 mi N Hermosa, Pennington County (SDMT); on the Pine Ridge Indian Res- ervation in nearby Shannon County; and at several localities in Harding County, north of the study area. Specimens of Onychomijs leuco^^asier from the Moor- craft area. Crook Co., Wyoming (just west of the Hills) are intergrades be- tween O. I. arcticeps and O. I. missouri- emis, but are assigned to the latter sub- species (Long, 1965:638). The northern grasshopper mouse may occur in the grasslands of the Red Valley "racetrack" in the Black Hills.
Lagurus curtatus levidensis (Gold- man, 1941 ) . — The sagebrush \'ole occurs west and southwest of the Black Hills in Campbell, Albany, and Laramie coun- ties, Wyoming (Long, 1965:660), north- west of the Hills in Carter and Powder River counties, Montana (Hoffmann et al, 1969:592), and northeast of the Hills in Williams and Morton counties. North Dakota (Hall and Cockrum, 1953:455). The foothills and canyons of southwest-
ern Custer County, South Dakota, and adjacent Weston County, Wyoming, are dominated by sagebrush and mountain mahogany that should provide suitable habitat for Lagunis. Although this mi- crotine has not been taken there as yet, the occurrence of preferred habitat in close proximity to the present known range of the species suggests that the sagebrush vole may inhabit the south- western edge of the Black Hills.
Vulpes velox hebes Merriam, 1902. — Of the swift fox, Grinnell (1875:80) wrote that "this pretty little fox is abun- dant everywhere on the plains," and Hoffman (1877:96) indicated that it was to be found in the xicinitv of the Black Hills. Much reduced by hunting, trapping, and poisoning pressures, this fox presumably could still survive in the Wind Cave National Park area. The species has been reported from Harding County in South Dakota (Visher, 1914: 90), southeastern Wyoming (Long, 1965:681), and northwestern Nebraska (Gary, 1902:67).
Urocyon cinereoargenteus ocythous
Bangs, 1899.— Long (1965:682) noted that the westernmost occurrence of this subspecies is represented b>' a large adult female from Owens, Weston Co., Wyoming (USNM 107892); howexer, I have not examined this specimen. Bailey (1927:166) indicated that the gray fox occurred along the northern border of the Black Hills, and an adult male was taken near Deer Ears Butte, Butte Co., South Dakota, in 1961 (Jones and Hen- derson, 1963:283). The "silver gray foxes" reported as being "numerous" in the Black Hills area by Dodge (1876: 123) may ha\e belonged to this species.
Spilogale putorius interrupta (Rafi- nesque, 1820). — Unfortunately, there seem to be no specimens of the spotted skunk available from the Black Hills region, even though this species un- doubtedly occurs there. Merritt Gary related to A. H. Howell (1906:7) that several spotted skunks were killed in the Black Hills, near Elk Mountain, Custer
TURNER: MAMMALS OF THE BLACK HILLS
153
Count)-, in 1902-03. In an unpublislied manuscript, Gary indicated that this spe- cies also occurred "near Newcastle" (Long, 1965:702). W. H. Over indi- cated in a letter to X. Schantz (1953: 125) that "I found a skull of this little mammal in a cave in the west slope of the Black Hills." E. R. Hall showed photographs of two spotted skunks, taken at the same spot in eastern Wyo- ming, to R. G. Van Gelder (1959:272); one displaxed the pattern of S. p. inter- nipta, and the other resembled S. p. <^rcicilis. Thus, these two kinds of spotted skunks mav not interbreed (see also Mead, 1968:389). It is most desirable tliat specimens be obtained from the Black Hills in the near future in order to shed light upon this taxonomic and biological problem. Van Gelder (1959: 251 ) included the Black Hills within the range of S. p. inferrupta.
Martes americana vulpina (Rafines- que, 1819). — Mr. J. Johnson captured a pine marten (USNM 249298) near Pringle, Custer Co., South Dakota on 4 January 1930. Whether this specimen represents an escaped pe*" or was actually indigenous to the Black Hills fauna re- mains uncertain. The species may have occurred in the region in the early 1800's; Hoffman (1877:96) reported seeing sev- eral specimens about 8 mi W Grand River Indian Agency, presently in Car- son County, South Dakota, and martens are known from southeastern Wyoming (Long, 1965:690).
Martes pennanti pennanti (Erxleben, 1777). — The fisher has been taken in northwestern Wyoming (Long, 1965: 691), and early trappers reported seeing it frequently in the Red River Valley
of North Dakota. It is possible that this species inhabited the Black Hills prior to the advent of European man in the region.
Gulo gulo luscus (Linnaeus, 1758). — The wolverine occurred occasionally in the Black Hills in the mid-nineteenth century (Baird, 1858:184; Coues, 1877: 49). The most recently recorded speci- men for South Dakota is a male killed by Mr. Kenneth Long on a sheep ranch near (south of) Timber Lake, Dewey County, on 10 April 1962 (Lee, 1962:21). Wolverines also have been recorded from northwestern Wyoming (Long, 1965:698) and western Nebraska (Jones, 1964:284).
Lutra canadensis canadensis (Schre- ber, 1776). — The otter once inhabited at least the major river systems on the Great Plains, and Grinnell (1875:80) speculated that it occurred in the Black Hills. Hoffman (1877:96) encountered occasional skins from west of the Grand River Agency.
Dama dama (Linnaeus, 1758). — The fallow deer is a cervid that has been in- troduced from Europe. A herd of fallow deer range over the Annenburg Ranch west of Spearfish, in Crook County, Wy- oming. A buck was shot near Hot Springs, Fall River County, in the au- tumn of 1955 by Hans Palmgreen, and another was killed near Piedmont in 1943 (South Dakota Conservation Di- gest, 25:16, July 1956). The irregularity with which these animals have been taken in the Black Hills suggests that the fallow deer has not become firmly established in the region, and that these specimens represent an occasional stray from the Annenburg Ranch.
FACTORS LNFLUENCING DISTRIBUTION AND SPECIATION
MAMMALIAN DISTRIBUTIONAL PATTERNS
Because the Black Hills represent a mountainous island habitat surrounded by grass, it is difficult to satisfactorily
place the Hills within previously pro- posed biotic distributional areas. Fur- thermore, the diverse topography and heterogeneous origins of the mammalian fauna in the Hills (see below) increase the difficulties. Nonetheless, the Life-
154
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
zone concept of C. H. Merriam (1899) has some merit, in a general way, when applied to the Black Hills. Because life- zones are based on temperature, the zonal boundaries correspond closely with isotherms. Kendeigh (1954) has re- viewed criticisms of this concept.
Transition Life-zone. — A warmth- adapted biota occurs on the arid prairie uplands and semiarid Red Valley that surround the Black Hills, in the decidu- ous riparian communities that cross these grasslands, and along the basal slopes of the foothills. Steppe-associated mammals (desert cottontail, black-tailed prairie dog, olive-backed pocket mouse, hispid pocket mouse, Ord's kangaroo rat, prairie \'ole, black-footed ferret, prong- horn, and bison) and riparian-associated mammals (eastern cottontail, fox squir- rel, and white-footed mouse) are fairly limited to the Transition Life-zone in the Black Hills region.
Canadian Life-zone. — A cold-adapted biota occupies the forested upper slopes of the foothills and the "boreal-cap" (Limestone Plateau, Central Basin, and Bear Lodge Mountains) of the Black Hills. Se\eral boreal or montane "iso- lates" are more or less restricted to this life-zone (e.g., red squirrel, northern Hying squirrel, yellow-bellied marmot, red-backed vole, and to an unknown ex- tent, the ermine). Other species, such as the least chipmunk, northern pocket gopher, and long-tailed weasel have en- demic subspecies in the Canadian Life- zone of the Black Hills, whereas the Transition Life-zone is inhabited by a different subspecies or represents a broad zone of intcrgradation between races. Some kinds of mammals (the masked shrew, Nuttall's cottontail, bushy-tailed woodrat, long-tailed vole, and meadow jumping mouse) are far more abundant on the "boreal-cap," than in the Transition Life-zone, where they are represented by much lower popula- tion densities. Many species of mammals are not confined to either life-zone, but abide in suitable habitats in each (the
white-tailed jackrabbit, thirteen- lined ground squirrel, deer mouse, and mea- dow vole) or range freely throughout both (large and mobile ungulates and carni\ores, xolant bats, and semiaquatic furbearers ) .
Long (1965:726-729) described sev- eral faunal areas in Wyoming, based on patterns of geographic distribution of mammals. In his system, the biota of the Black Hills is associated with the so- called Great Plains Faunal Area. Its divisions of immediate interest here are briefly discussed below.
The broad arid region that isolates the Black Hills from other forested mountains in northern and central Wy- oming comprises the Po\\xler River Valley-Bighorn Basin Faunal Di\ision. The long-tailed vole and Nuttal's cotton- tail range westward out of the Hills, along streamside environs in this region. The Cheyenne Plains Faunal Division isolates the Hills from the Laramie Mountains and other southern mountain ranges in Wyoming. A small tract of this unit is in contact ^^'ith the extreme southwestern corner of the Black Hills, projecting between the pre\ious and following faunal divisions, and support- ing typical plains species. The substrate of the Sand Hills Faunal Division is composed of sands or fine sandy loams that extend westward from the Sand Hills of Nebraska, along the South Da- kota-Wyoming border (Sand Hill Rego- sol Soil Subassociation), and is inhabited by various heteromyids and other are- nicolous mammals.
The Black Hills Faunal Division is delineated by the forested portion of the Hills and is inhabited by many mammals of boreal or montane affinities. The first three faunal di\isions, and the lower tracts of the fourth, are assignable to the Transition Life-zone, whereas most of the fourth faunal division is applicable to the Canadian Life-zone; representa- tive distributions of mammals in these life-zones were enumerated above.
TURNER: MAMMALS OF THE BLACK HILLS
155
ORIGIN OF THE RECENT
MAMMALIAN FAUNA
OF THE BLACK HILLS
The present biogeographic analysis of the Black Hills mammalian fauna is based on the distributions of 59 indige- nous species. These include four species (the wapiti, pronghorn, bison, and mountain sheep) that pre\iousl\- were extirpated and then reintroduced by
man, and one (the fox scjuirrel) that undoubtedly was introduced but prob- ably also reached the Black Hills along natural routes of dispersal. In addition, 11 species of uncertain status are in- corporated as a component of the Black Hills fauna; these .species either were re- ported from the Hills in early literature, or specimens ha\'e been recently ac- (juired immediately adjacent to the Hills (Table 29). Three .species (the Norway
Table 29. — Mammals of the Black Hills, listed by faiinal units as discussed in the text. An asterisk denotes species of uncertain status that i^roliably occur (or once did occur) in the
Black Hills region.
Widespread Species (26) Myotis leihii Mijotis hicifugus Lasionyctcris noctivagans Eptcsiciis fiiscus Lasiiinis cineretis Castor canadensis Peromyscus maniculatus Ondatra zihethicus Ercthizon dorsatum Canis latrans Canis lupus Vidpcs vulpes Ursus amcricanus Ursus arctos Procyon lotor Mustcla frcnata Miistela vison Taxidca taxtis Mcplntis mej)hitis Felis concolor Lynx rtiftis Cere us canadensis Odocoileus hemionus Odocoileus virginianus Antilocapra americana Bison bison
Steppe Species (10)
Lepus townsendii
Spermophilus tridecemlineatus
Cynomys ludovicianus
Perognath us fasciatus
Perognathus hispidus ° Reithrodontomys montanus
Microtus ochrogasier ''Vulpes velox
Mustela ni gripes "Spilogale putorius
(subspecies intcrrupta)
Great Basin Species ( 1 )
Eutaniias minimus (karyotype B)
Sonoran Species (7)
Myotis thysanodes
Sylvilagus auduhonii "'Lepus californicus "'Spermophilus spilosoma
Dipodomys ordii
Reithrodontomys mcgalotis "Onychomys leucogaster
Deciduous Forest Species ( 6 ) Myotis keenii Lasiurus borealis Sylvilagus floridantis Sciurus niger Peromyscus leucopus " Urocyon cinereoargenteus
Boreomontane Species (10) Sorcx cinereus Tamiasciurus hudsonicus Glaucomys sabrinus Clethrionomys gapperi Microtus pennsylvanicus Zapus hudsonius
"'Martes americana Mustela erminea
"Gulo gulo Lynx canadensis
Cordilleran Species (10)
* Sorcx nanus
"'Myotis evotis Myotis volans Plecotus townsendii Sylvilagus nuttaUii Marmota flaviventris Thomomys talpoides Neotoma cincrea Microtus longicaudus Ovis canadensis
156
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
rat, house mouse, and mountain goat) that were introduced from outside of the study area are not treated in this analy- sis.
Hoffmann and Jones ( 1970 ) recently presented a comprehensi\'e biogeo- graphic analysis of the Recent mam- malian fauna of the Northern Great Plains. Because this thorough study en- compasses the Black Hills region, much of the following account is based on the substance of their report. I also have drawn from my own summary of the vegetation and affinities of the Black Hills flora ( see pages 26 to 38 and Table 2).
Steppe Species. — Ten species (14.3%) of the Black Hills mammals have evolved in response to the semiarid to arid en- vironment of the interior North Ameri- can grasslands and are closely allied with the Great Plains biota. The distri- butions of Lepiis toicnsendu (listed with both the Great Basin and Campestrian faunal units by Armstrong, 1971), Spermophilus tridecemUneotus, Cijno- mys hidovicionus, Feroii,naihus fcisciatus, Microtus ocliro^o.ster, Vulpes velox and Mustela nigripes broadly oxerlap the Black Hills region, whereas distributions of Perognathus hispidus and Reithrodon- tomys montamis approach their north- westernmost limits in the Hills area, and Spilogale putorius (subspecies inier- rupta ) reaches its westernmost limit. The latter may be distinct from the Great Basin race, S. p. gracilis, at the specific level (Van Gelder, 1959:272; Mead, 1968:389). Two additional kinds {An- tilocapra americana and Bison bison) are intimately associated with the prairie grasslands but range beyond the central steppes and are treated with other eury- topic species. Likewise, Leptis californi- cus, Reithrodontomys megahtis, and Onychomys leucogaster are steppe-asso- ciated mammals but have a much more extensive distribution in the Southwest and adjacent Mexico.
During the maximal Wisconsin gla- ciation, much of the Northern Great Plains was covered by a boreal forest
and members of the current plains biota occupied steppe or savanna conditions to the south. Pollen profiles and the pres- ence of relict populations of Microtus ochrogaster {M. hidovicianus in western Louisiana and eastern Texas) and Cyno- mys hidovicianus (C. mexicanus in northern Mexico) are evidence of the southerly displacement of the Great Plains biota to suitable refugia in the Full-glacial period (Hoffmann and Jones, 1970:366). Although some re- stricted grassland may have occurred in the arid rain shadow of the Rocky Moun- tains in Late-glacial times, it was not until a shift in atmospheric circulation and ensuing climatic e\cnts occurred, in the Pre-Boreal and Boreal periods, that actual replacement of the boreal forest by grassland commenced on the Plains. Concomitantly, steppe-associated mammals dispersed northward, south of the ice front, and reinhabited the North- ern Great Plains and Black Hills region.
As the plains climate became increas- ingly warmer and drier, conditions fa- vored eastward extension of the grass- lands biota. Guilday et al. (1964) re- corded two prairie species, Spermophilus tridecemJineatus and Pedioecetes phasia- neUus (the sharp-tailed grouse), in east- ern Pennsylvania in the Late-glacial period. Mid-post-glacial time also is the only period that Spilogale putorius in- terrupta ranged east of the Mississippi River, into western Illinois (Parmalee and Hoffmeister, 1957). Shifts in the plains biota presumably occurred many times and to varying degrees, but most recent authors fa\'or the Atlantic period as the time of maximal eastward pene- tration of the grassland biota. In the Sub-Boreal period, the steppes retreated to their present limits.
Differential tolerance to high tempera- tures and accompanying heat stress may have dictated the post-glacial moxements of some members of the plains biota. For example, Microtus ochrogaster is much more tolerant of xeric conditions than is its boreomontane congener, xA/. pennsylvanicus; yet, populations of M.
TURNER: MAMMALS OF THE BLACK HILLS
157
ochrogaster ha\c decreased drastically in recent droughts ( Wooster, 1935, 1939; Gier, 1967). The easternmost margins of the plains-grasslands presumabK' would be somewhat more mesie, and the eastward shift in A/. ocJiwgaster dining drier post-glacial times may haxe been in response to decreased evaporative stress in such areas. Con\'ersely, the oc- currence of Pero!j.natlius Jiispidus in east- ern Missouri in Late-glacial or early Holocene time (Oesch, 1967) may be credited to the ability of this heteromyid to tolerate extremely cold temperatures (Jones, 1964:173). As a final example of temperature-influenced shifts in dis- tribution, the present northward retreat of Lepiis tounsendii from the southern part of its range and its current expan- sion to the north and northeast may be correlated with the gradual warming trend now in progress on the North American continent (op. cif.: 113).
Other factors influencing past distri- butions of steppe mammals include the presence of efl^ecti\'e barriers to dispersal and the availability of food sources. The Missouri River limited the eastward ex- pansion of P. hisvidus and Reithrodonto- )nys montanus. Past and present ranges of Musiela nifiripes correspond closely to those of Cynomys ludovicianns. There are some exceptions to the last men- tioned example. For instance, M. ni- iiripes is represented in late Pleistocene deposits in Jaguar Cave in Idaho, and in Orr Cave in Montana, but remains of Cynomys apparently were not present in either deposit (Guilday and Adam, 1967:30).
Sonoran Species. — Seven kinds of mammals (10.0%) present in the Black Hills are typical of the Chihuahuan- Sonoran Region of southwestern United States and adjacent Mexican Plateau. Like the steppe species, these mammals evidently originated under the dictate of an arid climatic regime, and subse- quently invaded the Hills region in times of post-glacial warmth and dryness, prol^ably in the late Boreal and early Atlantic periods. Most of these species
are associated with the interior grass- lands but have extensive distributions beyond the central steppes. Lepu.s cali- foniicu.s, Di])odomys ordii, Reithrodonto- mys me'^alotis and Onychomys leuco- gcister also range into the Great Basin, as does Myotic thysanodes. The ranges of Syhilagus- auduhonii and Spennophihis spdosovui are more widespread west of the Black Hills, but rarely exceed 40° latitude in the far west. Leptis ccdiforni- cus, R. megalotk, and O. leticogasier also are apportioned far east of the Hills, as was S. auduhonii during chronic droughts of a few decades ago (Hoff^- mann and Jones, 1970:382).
Both L. californicus and S. spilosoma approximate their northernmost limits on the Great Plains along the south- eastern margin of the Black Hills. The former species has extended northward into South Dakota within historic time, whereas the latter has not been taken in the Hills since the period of 1857-58.
Myotis thysanodes is represented by an endemic population in the Black Hills region (subspecies polia.sapen.sis). Either a few individuals managed to cross intervening barriers to initiate a disjunct colony, or the population in the Hills is a relict that resulted from the fragmentation of a formerly more wide- spread distribution.
Great Rasin Species. — Although many members of other faunal units range into the Great Basin (Armstrong, 1971, listed Myotis volan.s, Lepus townsendii, TJiomoinys talpoides and Antdocapra americana as components of this faunal unit), only one species (1.4%) of the Black Hills mammalian fauna presum- ably originated there. Hoffmann and Jones (1970:383-385) proposed a Great Basin-Great Plains pattern of differen- tiation for Eutamias niinimus, and indi- cated that subspeciation of Thomomys talpoides and Spdogale putorius also may fit this general pattern. Definitive evidence concerning differentiation of the latter two species is lacking as yet.
Recent analysis of chromosomal structure of Eutamias minimus (Sutton
158
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
and Nadler, 1969:526) indicates diat boreal forest races ( horeaJis and neglec- tus), Rocky Mountain subspecies (ari- zonemis and consohrinus), the Laramie Mountains race (operarius), and two secondarily steppe-adapted subspecies (niininuis of the Wyoming Basin and canji of the San Luis Valley) possess karyotype "A." The Great Basin races {scrutator and pictus). Northern Great Plains subspecies (palliclus and cacode- ?nus), and two montane populations (confinis in the Bighorn Mountains and silvatictis in the Black Hills) possess karyotype "B." The absence of hetero- morphic chromosomes is evidence that subspecies with these two different kary- otypes may in fact represent sibling species (Sutton and Nadler, 1969).
During maximal glaciation, the an- cestral stock of the Great Plains races of Eutamias minimus evidently became disjunct from topical boreal representa- tives. Isolated by a barrier of coniferous forest that intervened between the Great Basin and Wyoming Basin, the ancestral stock e\'ol\'ed in response to arid steppe and semidesert enxirons that were asso- ciated with sagebrush-grass communi- ties, as well as coniferous ^^'oodlands. The interposing coniferous forest barrier pre- sumably was intermittent, and formed in response to climatic conditions that fa\'ored each continental and Cordilleran glaciation; thus, it was fashioned by either southerly displacement of boreal forests or by descent of montane forest. For example, regional montane biotic zones throughout the cordillera were de- pressed vertically 4000-4500 feet, con- current with a decrease of 16-17 F in summer temperature in the Full-glacial period (Richmond, 1965:228, and oth- ers). A similar descent of the lower tree- line evidently occurred in the Sub-Boreal and possibly other periods.
As the boreal forest retreated north- ward and the montane woodland as- cended upslope, a relatively low non- forested pass opened across the Conti- nental Dixide between the Great Basin and Wyoming Basin. Thus the climate
of the early Holocene, or possibly of prior interglacial periods, permitted movement of Eutamias minimus across the W\'oming Basin, onto the Northern Great Plains. A shift of the upper-air antic>X'lonic eddy northeastward from the Great Basin in the Sub-Atlantic period ( Bryson and Wendland, 1967 ) may have influenced dispersal of the biota at a later time. The boreal forest chipmunks pre\iously in the plains region, theoreti- cally retreated upslope into montane en- vironments, coincident with retreat of boreal elements from the prairie. Ap- parently freed from intense competition, the invaders from the Great Basin then adapted to the central steppes environ- ment.
That Eutamias minimus confinis and E. m. silvaticus appear to have been derived from adjacent plains popula- tions rather than from boreal forest or Rocky Mountain ancestors is indicati\'e of several periods of isolation as de- scribed above. Otherwise, the montane derivatives of the plains populations would ha\e met extremely intense com- petition from residual boreal chipmunks, already occupying the Bighorn Moun- tains and Black Hills. Probably, boreal- adapted races would have a competitive advantage in a montane environment over those that were steppe-adapted. Under what conditions steppe-adapted races would find these montane areas free of boreal-adapted chipmunks, or would become competitively superior to the latter, remains problematical.
Deciduous Forest Species. — Six spe- cies (8.6%) of the Black Hills mammalian fauna have affinities with eastern hard- wood forests, mesic grasslands, or prairie riparian communities. Most of these mammals reach the westernmost limits of their continuous range in the xicinity of the Hills.
Mijotis keenii (subspecies septen- trionalis) and Sciurus niger are restricted to temperate eastern North America, whereas Lasiurus horcalis (a seasonal migrant in the Hills region), Sylvilagus jioridanus, Feromijscus leucopus and
TURNER: MAMMALS OF THE BEACK IIILES
159
Urocijon cinereoargenteiis aic> widely distributed in the temperate eastern and subtropical southern portions of the con- tinent. The last nientioncxl sp(>ci(\s orig- inated in the Neotropical Region, but invaded the Northern Great Plains from the east (Hoffmann and Jones, 1970: 377 ) . Whether Mijotis keenii is an actual "relict" or a seasonal migrant to the Black Hills is still in question. Peroniys- cus leucopus presently occurs on the Northern Great Plains in scattered lo- calities that are relatively isolated from their respective contiguous ranges (see Figs. 15 and 16, Hoffmann and Jones, 1970:38(8). The currently disjunct popu- lations of P. leucopus may represent iso- lated segments of a formerly more or less continuous and interbreeding population that dispersed northwestward behind the retreating ice sheet and was subse- (juently extirpated in the more xeric cli- matic regimes. Alternatively, they may typify the disruption of a continuous dendritic distribution that paralleled mesic drainage systems westward across the prairies. The latter explanation is favored by most recent authors. Cur- rently, gene-flow between these limited populations is reduced or absent, and each presumably is adapting independ- ently to its immediate en\'ironmental conditions, as evidenced by the rela- tively small and dark mice on the Black Hills.
Elements of the eastern deciduous forest replaced boreal forest species along the eastern periphery of the North- ern Great Plains in Pre-Boreal times. Members of this faunal unit presumably then dispersed westward along wooded tributaries of the Missouri River system, only to be excluded from the plains (or isolated in suitable refugia) in the sub- sequent arid Atlantic time. With ameli- oration of climatic conditions in the more mesic Sub-Atlantic and Neo-At- lantic periods, eastern species could dis- perse westward along gallery forests once more. Several species (Sciurus niger and Urocyon cinereoargenteus) have extended their ranges westward to
the Black Hills area within historic time.
WUles-pread Species. — Twenty-six species (37.1%) of the Black Hills mam- mals are eurytopic, with no apparent faunal affinities in relation to the Hills region. These kinds are present in the study area because they typically are extremely mobile. They are euryecious with a wide range of tolerance for vari- ous environmental factors, or are sten- oecious but encounter specific environ- mental requisites in several segregated habitats throughout the region.
Volant bats, and large ungulates and carnivores, are exceedingly mobile, rang- ing over vast areas that may be com- posed of several different environs. Of the eurytopic chiropteran fauna, Myotis leibii is restricted to temperate North America, whereas M. lucifugiis and Lasionycteris noctivagans also range over boreal portions of the continent, and Eptesicus fiisctis and Lariurus cine- reiis additionally are distributed to the Neotropical Region. Two of these spe- cies (L. noctivagans and L. cinereus) are seasonal migrants in the Black Hills.
Of the large game mammals and fur- bearers whose distributions overlap the Black Hills, five are primarily temperate kinds ( Taxidea taxiis, Lynx rufus, Odo- coileus hemionus, Antilocapra americana and Bison bison), three are mainly tem- perate-boreal taxa (Vidpes vulpes, Me- phitis mephitis and Cervus canadensis), three are temperate-boreal-subtropical types (Canis hipus, Ursus americanus, and U. arctos) and five species (Canis latrons, Procyon lotor, Miistela frenata, Felis concohr, and Odocoileus virgini- ana) range from temperate regions into both boreal and Neotropical areas. Three of these mammals ( U. americanus, P. lotor, and O. virginiana) are primarily associated with woodlands throughout their ranges, whereas four others (T. taxiis, O. hemionus, A. americana, and B. bison) are associated for the most part with nonforested environs. The re- maining nine species occur in both major habitats.
Three stenoecious aquatic or semi-
160
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
aquatic mammals are included in the eurytopic faunal unit, Castor canadensis. Ondatra zihethicus, and Mustela vison. These are temperate-boreal species with specialized niche requirements that are fulfilled by a variety of aquatic habitats throughout their range.
Among widespread terrestrial ro- dents, Erethizon dorsaitim is a temper- ate-boreal species that usually is asso- ciated with coniferous trees, but has an extensive distribution beyond such for- ests. Peromyscus manlculatus best ex- emplifies a euryecious taxon. This species consists of a long series of interbreeding, or potentially interbreeding, populations distributed from northern boreal North America southward to Oaxaca, Mexico. It is ubiquitous, occupying a remarkable variety of habitats and, being quite re- sponsive to local environmental selective pressures, has formed numerous eco- types throughout its range.
Boreomontane Species. — Ten species (14.3%) of the mammals inhabiting the Black Hills are distributed both in the northern boreal forest and in the conif- erous forest of western montane areas. Mammals of the boreomontane faunal unit for the most part range much far- ther north than do members of the cor- dilleran faunal unit; southern limits of distribution also average farther north for the former than for the latter.
Boreal forest, displaced southward by continental glaciation, and montane forest, displaced downslope, eastward and southward by Cordilleran glacia- tion, probably intermixed along a broad front, maintaining a continuous conifer- ous woodland throughout the Powder River Valley and Cheyenne Rixer plains. The boreomontane biota in the Black Hills was thereby connected with that of the Bighorn and Laramie mountains until at least the Pre-Boreal period. Sev- eral forested corridors that provided avenues for dispersal of boreomontane mammals remained on escarpments and other topographic breaks well after the coniferous forest retreated upslope and
northward (Hoffmann and Jones, 1970: 386 and Fig. 1).
Relict populations of northern mam- mals that now are disjunct from other such populations, and from the main range of the species, are an indication of the widespread extirpation of boreal elements during the Atlantic period and other warm, dry climatic episodes. Only the hardiest colonies survived the xeric conditions, and these abided in the fa- vorable or tolerable cool and mesic pock- ets on the plains, or in montane regions.
Three boreomontane mammals ( Sorex cinereus, Microtus pennsylvanicus and Zapiis hudsonius) are widely distrib- uted across the Northern Great Plains in suitable riparian communities. The lat- ter species lacks a western montane distribution typical of other members of this faunal unit, and often occurs in populations that are isolated from oth- ers by broad expanses of inhospitable terrain.
A few species (Martes americana, Gulo gulo and Lynx canadensis) repre- sent northern kinds that presumably in- habited the Black Hills in the recent past, but ha\'e since become extinct there. Occasionally, some representatives of these species may still wander into the Hills region.
The remaining four species (Tamia- sciurtis hudsonicus, Glmicomys sahrinns, Clethrionomys ii^apperi, and Mustela er- minea) represent "glacial relicts" that probably were isolated subsequent to the northerly retreat of the ice sheet. Mustela erminea is disjunct in the Black Hills to an unknown extent; it is present in the Laramie Mountains, but appar- ently absent in the Bighorn Mountains. The level of divergence of Clethriono- mys gapperi ])revicaudus (previously regarded as a distinct species) suggests that this \'o]e may ha\'e become disjunct earlier than did other "isolates," or may have diverged at a faster rate; the op- posite supposition may apply to Glau- comys sabrinus and M. erminea (see the following section on Speciation). Cock- rum and Fitch (1952) indicated that
TURNER: MAMMALS OK THE BLACK HILLS
161
C. <!,. l)revicau(lus is closely related to C. g. g,alei, and thus may have cordil- leran-montanc> affinities. However, until more evidence is a\ ailable, my comments here are based on the more northern distribution of the species as a unit.
The subspecies of T. hudsonicus in the Black Hills (cJakotensis) appears to be more closely related to the reddish- colored boreal forest populations than to the darker-colored Rocky Mountain races (Hoffmann and Jones, 1970:372). Reddish-colored squirrels in the north- ern Laramie Mountains, southeastern Montana, northwestern South Dakota, and other outlying areas probably are remnants of the former population that remain distributed along past corridors and dispersal routes. Assuming that da- kotensis originated in response to selec- ti\e pressures of the Black Hills environ- ment, the race either enjoyed a broader distribution in early Holocene time, or has since dispersed outward from the Hills via pine-filled canyon systems and conifer-dominated escarpments.
Cordilleran Species. — Ten species (14.3%) of the Black Hills mammalian fauna represent elements that were dis- placed into the region by Cordilleran ice sheets. These mammals evidently became stranded after retreating upslope in response to increasing aridity in post- glacial time. All but three species attain their easternmost limits on the Northern Great Plains in the Black Hills— the range of Neotoma cinerea and Ov)is cana- densis extend slightly beyond, terminat- ing in western South Dakota, and Thom- omijs tolpoides is broadly distributed eastward to approximately 98 longi- tude. The plains and montane races of the latter species may have a pattern of differentiation similar to that of Eutani- ias minimus.
Myotis evotis, M. vohns, Plecotus toicnsendii, 'Neotoma cinerea and Ovis canadensis generally are associated with montane forests, rocky outcrops, or bad- lands topography in western North America, whereas Sorex nanus usually is associated with subalpine or alpine rock
slides in the central-western United States. Sylvilaii,iis nuttallii and Microtus loniiicaiidus hav(> relatively broad niches, dispersing westward across the Powder Ri\'er Valley plains in association with riparian communities. Of the cordilleran faunal unit, only Marniota flaviventris represents a distinctly disjunct popula- tion. Isolation of this marmot in the Black Hills presumably occurred in the manner assumed for other "glacial re- licts."
Several taxa present in the Bighorn and Laramie mountains either failed to disperse to the Black Hills, or were extir- pated there during xeric climatic regimes (Hoffmann and Jones, 1970:376 and Table 2). This fact, in conjunction with the presence of other taxa in the Big- horn Mountains that are absent in the Laramie Mountains, and vice versa, sug- gests a filtering effect of past barriers, or differential survival of species reaching these montane environments, or both.
Using the distribution maps of Long (1965), the number of subspecies of mammals in each of the three indicated montane environs was ascertained. Twenty-five taxa occur in all three ranges, and 19 kinds are common to the Black Hills and Laramie Mountains, but do not abide in the Bighorns. Four sub- species are shared by the Black Hills and Bighorn Mountains, but not by the Lara- mie Mountains. Eleven races are com- mon to the Bighorn and Laramie moun- tains, but do not occur in the Black Hills. Sixteen kinds inhabit the Black Hills, but are absent from the other two ranges. Eleven subspecies occur in the Bighorn Mountains, but not in the Black Hills or Laramie Mountains, and 13 subspecies live in the Laramie Mountains, but not in the Black Hills or Bighorns. Three species (Etitamias minimus, Marmota fiaviventris, and Thomomijs talpoides) have diversified into separate subspecies in each of the aforementioned mountain- ous regions.
Employing these data, a percentage index of faunal resemblance (Simpson, 1943) can be calculated for each pair
162
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
of montane areas. Such indices are not absolute, but are meaningful only when comparing neighboring faunas, thereby yielding a relati\e measure of similarity (Udvardy, 1969:273). The larger the percentage index, the more closely the faunal pairs resemble each other. Com- puted indices between the various pairs are as follow: Black Hills and Bighorn Mountains, 56.9; Black Hills and Lara- mie Mountains, 68.7; Bighorn and Lara- mie mountains, 70.6.
Biogeographic analysis of affinities of the mammals in the Bighorn and Lara- mie mountains were undertaken in a manner similar to that used in the analy- sis of the Black Hills biota. Percentages of the mammalian fauna attributable to the respective faunal units are given for the Black Hills, followed sequentially by that of the Bighorn Mountains and Lara- mie Mountains as follow: widespread species, 37.1, 41.2, 36.8; steppe species, 14.3, 9.8, 11.8; Great Basin species, 1.4, 2.0, 2.9; Sonoran species, 10.0, 7.8, 8.8; deciduous forest species, 8.6, 0.0, 4.4; boreomontane species, 14.3, 13.7, 11.8; cordilleran species, 14.3, 25.5, 23.5.
Indices of faunal resemblance and biogeographic analyses both suggest that coniferous forest corridors between the Bighorn Mountains and Black Hills were indirect — \'ia the northern Laramie ranges (Casper and Haystacks), the Hartville Uplift, and the Pine Ridge escarpment (Hoffmann and Jones, 1970: 386 ) . This would account for the greater faunal similarities between the Bighorn and Laramie mountains, and between the Laramie Mountains and Black Hills. The relatively high percentage of cor- dilleran mammals in the two former montane environs, as compared to the Hills, further implies that the indirect connection was of greater duration, or was a much better dispersal route, or both, than was a direct extension of coniferous forest eastward from the Big- horn Mountains to the Hills.
Whereas the mammalian faunas of the Bighorn and Laramie mountains are similar due to a predominance of cor-
dilleran kinds, the Laramie Mountains fauna resembles that of the Black Hills due to a greater presence of steppe, So- noran, and eastern species. The Black Hills and Bighorn Mountains, on the other hand, both exceed the Laramie Mountains in relative proportions of boreomontane and widespread mam- mals. The latter mountainous region is intermediate in ele\ation ( and perhaps other environmental factors) as com- pared to the former two montane areas. Another possible avenue of exchange between the Black Hills and Bighorn Mountains may ha\'e occurred to the north, via the wooded Rosebud-Tongue river breaks and conifer-clad Long Pine Hills of southeastern Montana and northwestern South Dakota.
SPECIATION AND GEOGRAPHIC VARIATION
BioticaUy-induced Selection Pres- sures.— Two pairs of related species {Mijotis evotis and M. thysanodes; Syl- vihgiis floridanus and S. nuttallii) are parapatric in the Black Hills region. Several other pairs of species {Sorex cinereiis and S. nanus; Sylvilagus audti- honii and S. floridanus; Reithrodonio- mys megalotis and R. montanus; Pero- myscus leucopus and P. maniculatus; Microtus longicaudiis and M. pennsyl- vanicus; M. ochrogaster and M. penn- sylvanicus; Mustela erminea and M. frenata; Odocodeus hemionus and O. virginiana) are known to be sympatric, but ecologically distinct to varying or unkno\\'n degrees. Specific distinctness of such species pairs strongly suggests extrinsic isolation in the past and subse- quent development of intrinsic isolating mechanisms in response to differential selection pressures. Indeed, with the exception of the eurytopic deer, each member of the aforementioned species pairs belongs to a separate faunal unit of different biogeographic affinity (see Table 29). Interaction among closely related species favors differential ex- ploitation of similar niches, thereby de-
TURNER: MAMMALS OF THE BLACK HILLS
163
creasing the intensity of interspecific competition.
Seemingly, selection pressures would be less stringent on those species that do not compete with congeners in the study area. However, interspecific and ecologi- cal interactions between various sym- patric genera ( Lepus and Sylvilagus; Eufamias and SpenuopJiihis; GJaucomys, Sciiinis and Tainiasciunis; Cletliriono- mys and Micwtiis), among various chi- ropterans, and among several heteromy- ids may favor partitioning of similar niches in the Hills region due to these biotically-induced selection pressures.
EnvironmenfaUy-indiiced Selection Pressures. — Many kinds of mammals have differentiated into more or less dis- tinct subspecies as the result of selection pressures engendered by the Black Hills environment. Four of these subspecies ( Myotis thysanodes pahasapensis, Ta- miasciunis hudsonicus dacotetisis, Mar- nwfa flaviventris dacoto, and Clethriono- mys gapperi brevicaudus) currently are disjunct and continue to adapt inde- pendently. Three others (Eutomias mininuis silvaticiis, Thomomys talpoides nelndosns and Miistelo frenata olleni) are represented on the surrounding plains by different subspecies (E. m. pall id us, T. t. hidhtus and T. t. pierreic- olus, M. f. longicouda and M. f. neva- derisis), and are presumed to be in ge- netic contact with these surrounding populations. The presence of parapatric subspecies implies either that the two races formerly were isolated and di- verged (but not to the extent of devel- oping intrinsic isolating mechanisms) in response to different environments be- fore subsequent contact with the step cline, or that subspecific divergence oc- curred in situ due to strong selective pressures that formed separate clinally connected ecotypes. The extreme con- trast between the Black Hills and Great Plains in climate, topography, and bi- otic communities enhances the feasibility of the latter mechanism. The general pattern of \'ariation in other mammalian species (see following discussion) also
suggests in situ divergence of clinally connected ecotypes in the Black Hills region.
For the most part, the Black Hills are inhabited by races of mammals that also inhabit the encompassing plains. Continual exchange of hereditary ma- terials between the Hills and plains populations seems probable. Dichroma- tism (dark and pale phenotypes) and extreme individual variation in the Black Hills populations of Peromyscus manicu- latus- (Osgood, 1909:78; Dice, 1942:1- 10), and of Spernwphilus tridecemline- atus and Microtus longicaudus (Long, 1965:738) presumably are due to dis- ruption of the strong selection for darker coloration in the humid Hills by influx of alleles from paler forms on the arid plains.
An additional source of variability in mammals within the Hills is due to the great diversity in physiognomy, exposure of soils, and other general edaphic char- acteristics. Because small mammals are intimately associated with their habitats, selection pressures in one area may be quite different from those in another.
For example, the relatively dry, hard- wood-dominated drainage system of Big Spearfish Canyon and similar streambeds in the Bear Lodge Mountains contrast sharply with the cool, mesic Little Spear- fish Canyon that is characterized by lux- uriant moist meadows and spruce-bor- dered streams. As indicated in Figure 12, the general climatic regime of the north- ern Hills differs from that of the southern Hills. Additionally, the climatic and en- vironmental gradations from relatively mesic higher elevations, to semiarid foothills, to arid plains is much more gradual in the southern Hills as com- pared to the abrupt changes typical of the northern Hills (see page 112). The occurrence of Eutamias minimus polli- dus, rather than E. m. silvaticus in the extreme southern and southeastern Black Hills probably is a reflection of the grad- ual gradation of environmental factors, hence, of environmentally-induced selec- tion pressures. Other small mammals in
164
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
the southern portion of the Black Hills generally tend to be slightly paler than those of the northern area. In addition, the dorsal pelage of several mammals in the northwestern sections reflect red hues more intensely than those from other areas. Species that \'ary noticeably within the Black Hills include Eutamias minimus, Tamiasciurus hudsonicus, Tho- momys talpoides, Peromyscus manicu- latus, Neotoma cinerea and Microtus pennsylvaniciis.
The major effect of selection in the Black Hills as a unit seems to be in accord with Gloger's Rule, producing relatively dark and reddish-colored pop- ulations. The positive correlation in- dicated by Stebler (1939) and Dice (1942) between the color of dorsal pelage of mammals and the color of exposed soils in the Black Hills and on the Great Plains already has been dis- cussed (see page 99). Dice (1947:19), after undertaking several experiments that established significant selection by barn owls for deer mice contrasting in coloration with their background, con- cluded that ". . . natural selection can theoretically produce rapid evolution whenever a genetically variable popula- tion is exposed to its action." Popula- tions presumably respond to diff^erent environments through variation in gene frequencies, but natural selection tends to minimize variability, increasing the frequency of adaptive phenotypes and thus creating local ecotypes.
Clark (1938) indicated that the pale ochraceous-buff color of deer mice on the plains is a character dominant over the recessive darker coloration that fre- quently occurs in deer mice inhabiting montane areas. Seemingly, the environ- ment in the Black Hills favors a higher frequency of recessive dark alleles, but the gene pool is disrupted by the influx of dominant pale alleles from the sur- rounding plains populations, resulting in dichromatism within Hills popula- tions. If a similar genetic scheme may be assumed for the several isolated mam- mals in the Black Hills region, then ge-
netic drift may also play a role in in- creasing recessive homozygosity, thereby enhancing differentiation of these dis- junct populations.
The broad arid regions (Belle Fourche, Cheyenne, and Powder river plains) that isolate the Hills from other montane areas are inadequate barriers to some species. These rivers provide routes of immigration and emigration for Sorex cinereus, Sylvilagus floridanus (to the north, south and east), Sylvila- gus nuttoUii (to the west), Sciurus niger, Thomomys talpoides, Peromyscus leuco- pus, Microtus longicaudus, Microtus pennsylvanicus, Zapus liudsonius, and various semiaquatic mammals. Thus, genetic flow is probable along streams in these species, between riparian-resti'icted plains populations and those on the Black Hills. Long (1965:733) noted that among lagomorphs, the species most de- pendent on riparian habitats is darkest in color — Sylvilagus floridanus (dark), S. tiuttallii (intermediate), and S. audu- bonii (pale).
The degree of differentiation of dis- junct species in the Black Hills may be used as an indirect measure of the dura- tion of such isolation. Such indications are relative and may be modified greatly by unknown differential rates of evolu- tion of the species in question. For ex- ample, Clethrionomys gapperi hrevi- caudus differs from C. g. galei of the Rocky Mountains in 11 mensural param- eters ( see also Cockrum and Fitch, 1952 ) and previously was treated as a distinct species (Bailey, 1897:129). The remain- ing subspccific isolates (Myotis thysa- nodes, Tamiasciurus hudsonicus, and Marmota flaviventris) differ from the nearest conspecific populations mainly in color, or differ in mensural characters to a comparatively lesser extent than en- countered between the two subspecies of Clethrionomys gapperi. Myotis keenii, Glaucomys sahrinus, and Peromyscus leucopus also are isolated to differing degrees and currently appear to be di- verging under thc^ dictate of natural se- lection in the Black Hills. The Hills
TURNER: MAMMALS OF TJIE BLACK HILLS
165
population of Mustela erminea appears disjunct, but cnidcntly does not differ notabh' from members of this species in the Laramie Mountains. Hence, C. gap- jicri either became isolated in the Black Hills at an earlier time than did the other "glacial relicts," or has diverged at a faster rate due to a shorter genera- tion time. Conversely, those species that ha\e differentiated in the Hills region to a lesser extent either were isolated in a later time period or ha\e responded more slowly to selection pressure in- curred by the Black Hills environment. Manv kinds of mammals with broad distributions that oxerlap the Black Hills have not diverged subspecifically, but display some modification in coloration within the Hills. Populations of Spermo-
pliilus tridecemlineatus, Neotoma cin- erea, Peromyscus maniculatus, Microtus longicaudtis, Microtus penn.sylvanicus and Microtus ochrogaster all seem to be evolving darker color in the study area. On the other hand, representatives of Reitlirodontomys megalotis that occur in the Hills show no appreciable change in coloration when compared to those of the surrounding plains. Finally, tliree species (Sorex cinereus, Peromyscus moniculotus, and Zapus hudsonicus) vary clinally in coloration, and in exter- nal and cranial dimensions northwest- wardly across the Great Plains ( Dice, 1941:16; Robertson, 1971). With excep- tion of somewhat darker color, the Black Hills apparently do not interrupt the clinal nature of variation in these species.
SUMMARY
The Black Hills of western South Dakota and northeastern Wyoming con- stitute an elliptically-shaped, isolated mountainous region of approximately 4000 square miles that resulted from intermittent domal uplifts in Cretaceous, Miocene, and Pleistocene times. En- tirely surrounded by the non-glaciated Missouri Plateau section of the Northern Great Plains physiographic province, the Hills rise above the plains about 4000 feet on the east and some 3000 feet on the west; the highest point is 7242 feet above sea level. Most of the area lies within the drainages of the Cheyenne and Belle Fourche rivers. As here con- sidered, the Black Hills are delimited by the distribution of Jurassic shale and sandstone of the Sundance Formation. The region thus circumscribed includes the Black Hills proper, as well as the Bear Lodge Mountains and Devils Tower of Wyoming; these tracts are closely allied geographically, geologically, cli- matically, physiognomically, and biologi- cally. Ranges of mountains nearest to the Black Hills are the Laramie Moun- tains (to 10,272 ft) and Big Horn Moun- tains (to 13,165 ft), approximately 150
miles to the southwest and west, respec- tively, in Wyoming.
This report delimits and describes the Black Hills as a natural zoogeo- graphic unit, considers the autecology and general distribution of each mam- malian species in the Hills, discusses the geographic variation and inferred specia- tion of these mammals, and analyzes and interprets the probable biogeo- graphic affinities of various species in light of conjectured changes in late Pleistocene and Holocene environments. Six orders and 19 families, including 44 genera and 62 species, comprise the Black Hills mammalian fauna.
Biogeographical analysis of the Recent mammalian species of the Black Hills suggests the following faunal affinities: 37.1 percent mdespread, 14.3 percent steppe, 14.3 percent boreomontane, 14.3 percent cordilleran, 10.0 percent So- noran, 8.6 percent deciduous forest, and 1.4 percent Great Basin. Indices of faunal resemblance and biogeographic analyses suggest that coniferous forest corridors between the Bighorn Moun- tains and Black Hills in the Pleistocene
166
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
time were indirect; this accounts for the faunal similarities between the Bighorn and Laramie mountains, and between the Laramie Mountains and Black Hills. The relatively high percentage of cor- dilleran mammals in the two montane environs, as compared to the Hills, fur- ther implies that the indirect connection was of greater duration, or was a much better dispersal route ( or both ) than was a direct extension of coniferous forest eastward from the Bighorn Mountains to the Hills.
Interaction among closely related spe- cies favors differential exploitation of similar niches, thereby decreasing the intensity of interspecific competition. Ten pairs of related species in the Hills region either are parapatric or sym- patric but ecologically distinct to varying degrees. Specific distinctness of such species pairs strongly suggests extrinsic isolation in the past and subsequent development of intrinsic isolated mech- anisms in response to differential selec- tion pressures. Intergeneric competition also may favor partitioning of similar
niches due to these biotically-induced selection pressures.
Four endemic subspecies currently are disjunct from contiguous popula- tions, whereas three others are in genetic contact with different subspecies on the i surrounding plains. Such parapatric sub- 1 species probably di\'erged in situ due to local selection pressures that formed separate, clinally-connected ecotypes.
Most races in the Hills also inhabit the plains. Dichromatism in several spe- cies is due to disruption of selection for darker coloration in the humid Hills by influx of pale alleles from arid plains populations. Small mammals in the southern Hills tend to be slightly paler than those in the northwestern Hills, which reflect red hues more intensely than individuals from other areas.
Many eurytopic mammals have not diverged subspecifically in the Hills, but display modification in coloration; one species shows no apparent change. Three species vary clinally northwestwardly across the plains; the Black Hills do not interrupt these clines.
LITERATURE CITED
Allen, G. M.
1916. Bats of the genus Corynorhinus. Bull. Mus. Comp. Zool., 60:331-356.
1931. Type specimens of mammals in the Museum of Comparative Zoology. Bull. Mus. Comp. Zool., 71:227-289. Allen, H.
1864. Monograph of the bats of North America. Smithsonian Misc. Coll., 7:x.\iii+l-85.
1894. A monograph of the bats of North America. Bull. U. S. Nat. Mus., 43: ix-^1-198. Alle.v, J.A.
1877. History of the American bison, Bison americanus. Ann. Rept. U. S. Geol. Surv. Terr., 9:443-587.
1890. A review of some of the North Ameri- can ground squirrels of the genus Tamias. Bull. Amer. Mus. Nat. Hist., 3:45-116.
1894a. Descriptions of ten new North Ameri- can mammals, and remarks on others. Bull. Amer. Mus. Nat. Hist., 6:317- 333.
1894b. Descriptions of five new North Ameri- can mammals. Bull. Amer. Mus. Nat. Hist., 6:347-350.
1895a. List of mammals collected in the Black Hills region of South Dakota and western Kansas by Mr. Walter W. Granger, with field notes by the col- lector. Bull. Amer. Mus. Nat. Hist., 7:259-274.
1895b. Descriptions of new American mam- mals. BuU. Amer. Mus. Nat. Hist, 7:327-340.
1899. Descriptions of five new American rodents. Bull. Amer. Mus. Nat. Hist., 12:11-17. Andersen, K. W., and J. K. Jones, Jr.
1971. Manmials of northwestern South Da- kota. Univ. Kansas Publ., Mus. Nat. Hist., 19:361-393. Anderson, R. M.
1943. A prior name for tlie bean mouse re- vived. Canadian Field-Nat., 57:92. Anderson, S.
1956. Subspeciation in the meadow mouse, Microtus pennsijlvanicus, in Wyoming,
TURNER: MAMMALS OF THE BLACK HILLS
167
Colorado, and adjacent areas. Univ. Kansas Pulil., Mus. Nat. Hist., 9:85- 104.
Armstrong, D. M.
1971. Distribution of mammals of Colorado. Monog. Mus. Nat. Hist. Univ. Kansas, 3:x+415 pp.
Austin, M. E.
1965. Land resource regions and major land resource areas of the United States. Handbook U. S. Dept. Agric, 296:1- 82. Baerreis, D. a., and R. A. Bryson
1965. Climatic episodes and the dating of the Mississippian cultures. Wisconsin Archeol., 46:203-220.
Bailey, V.
1888. Mammals. Pp. 431-454, in Report on
some of the results of a trip through
parts of Minnesota and Dakota. Ann.
Rept. Dept. Agric. (for 1887), pp.
426-454. 1893. The prairie ground squirrels or sperm-
ophiles of the Mississippi Valley. Bull.
U. S. Dept. Agric, 4:1-71. 1897. Revision of the American voles of the
genus Evotomys. Proc. Biol. Soc.
Washington, 11:113-138. 1900. Revision of American voles of the
genus Microtiis. N. Amer. Fauna, 17:
1-88.
1907. Wolves in relation to stock, game, and the National Forest Reserves. Bull. U. S. Forest Serv., 72:1-31.
1908. Destruction of wolves and coyotes. Circ. U.S. Dept. Agric, 63:1-11.
1914. Eleven new species and subspecies of pocket gophers of the genus Thomo- niys. Proc. Biol. Soc. Wasliington, 27: 115-118.
1915. Revision of the pocket gophers of the genus Thomomijs. N. Amer. Fauna, 39:1-136.
1920. Identity of the bean mouse of Lewis
and Clark. Jour. Mamm., 1:70-72. 1924. Breeding, feeding, and other life hal>
its of meadow mice (Microtus). Jour.
Agric. Res., 27:523-536. 1927. A biological survey of North Dakota.
N. Amer. Fauna, 49:vi-f-l-226.
Baird, S. F.
1858. Mammals. In Reports of exploration and surveys . . . from the Mississippi River to the Pacific Ocean . . ., 8: xxi-xlviii-}- 1-757.
Baker, R. H., and J. S. Fixdley
1953. Sorex vagrans first reported from South Dakota. Jour. Mamm., 34:382.
Baker, R. H., and R. M. Haxoxs
1950. A new subspecies of snowshoe rabbit in Wyoming. Proc. Biol. Soc. Wash- ington, 63:63-64.
Banfield, a. W. F.
1962. A revision of the reindeer and caribou, genus Rangifer. Bull. Nat. Mus. Can- ada, 177:vi-f-l-137. Bangs, O.
1896. A review of the squirrels of eastern North America. Proc. Biol. Soc. Washington, 10:145-167. Barbour, E. H.
1931. The musk-oxen of Nebraska. Bull. Nebraska State Mus., 1:211-223. Benninghoff, W. S., and C. W. Hibbard
1961. Fossil pollen associated with a late- glacial woodland musk-ox in Michi- gan. Papers Michigan Aced. Sci., Arts, and Letters, 46:155-159. Berner, L. M.
1952. Antelope restoration. South Dakota Dept. Game, Fish and Parks, Pittman- Robertson Proj., 22-D-2:l-10.
1953a. Black clouds over our deer range. South Dakota Conserv. Dig., 20:4-5.
1953b. Deer management in the Black Hills of South Dakota. Part I. South Da- kota Conserv. Dig., 20:8-13, 16.
1953c. Deer management in the Black Hills of South Dakota. Part II. South Da- kota Conserv. Dig., 20:8-12, 16.
1954. Antelope seasons past and present. South Dakota Conserv. Dig., 21:2-3.
1955. History of antelope in South Dakota. Soutli Dakota Conserv. Dig., 22:2-3.
Bever, W.
1954. Changes in browse density and com- position on the Annie Creek and Hep- ler Range study areas from 1948 to 1953. South Dakota Dept. Game, Fish and Parks, Pittman-Robertson Proj., 12-R-11:1-12.
1955. Rocky Mountain goats, trapped and transplanted. South Dakota Conserv. Dig., 22:3-4, 10.
1957. The deer of South Dakota. South
Dakota Conserv. Dig., 24:4-5, 12. 1959. Deer, land, people. South Dakota Conserv. Dig., 26:2-8. Block, D. W.
1970. Lungworms of mountain goats in the Black Hills of South Dakota. M.S. thesis. South Dakota State Univ., 117 pp. Boddicker, M. L.
1970. Helminth parasites of wild ruminants in South Dakota. M.S. thesis. South Dakota State Univ., 146 pp. Boddicker, M. L., and E. J. Hugghins
1969. Helminths of big game mammals in South Dakota. Jour. Parasit., 55: 1067-1074. Boddicker, M. L., E. J. Hugghins, and A. H.
RlCHARDSON
1971. Parasites and pesticide residues of mountain goats in South Dakota. Jour. Wildlife Mgt., 35:94-103.
168
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Bole, B. P., Jr.
1935. Myotis thysanodes in South Dakota. Jour. Mamm., 16:147-148. Bole, B. P., Jr., and P. N. Moulthrop
1942. The Oliio recent mammal collection in the Cleveland Museum of Natural History. Sci. Publ. Cleveland Mus. Nat. Hist., 5:83-181. Brackett, a. G.
1881. The lynxes. Amer. Field, 16:413. Brady, D.
1955. Northern Hills deer wintered well. South Dakota Conserv. Dig., 22:13. Braun, E. L.
1947. Development of the deciduous forests of eastern North America. Ecol. Monogr., 17:211-219. 1951. Plant distribution in relation to the glacial boundary. Oliio Jour. Sci., 51:139-146. 1955. The phytogeography of unglaciated eastern United States and its inter- pretation. Bot. Rev., 21:297-375. Brown", L. N.
1967. Ecological distribution of six species of shrews and comparison of sampling methods in the central Rocky Moun- tains. Jour. Mamm., 48:616-623. Bryan, F. T.
1858. Reports of Francis T. Bryan . . ., Exec. Doc, U. S. Senate, 35th Cong., 1:455-481. Bryson, R. a., D. a. Baerreis, and W. W. Wexdlaxd
1970. The character of late-glacial and post- glacial climatic changes. Pp. 53-74, in Pleistocene and Recent environ- ments of the Central Great Plains ( W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp. Brysox, R. a., and W. M. Wexdland
1967. Tentative climatic patterns for some late-glacial and post-glacial climatic episodes in central North America. Pp. 271-298, in Life, land and water (W. J. Mayer-Oakes, ed.), Univ. Manitoba Press, Winnipeg, xvi-|-414 pp. Btjechxer, H. K.
1960. The bighorn sheep in the United States, its past, present, and future. Wildlife Monogr., 4:1-174.
Burt, W. H., and R. P. Grossenheider
1964. A field guide to the mammals. Hough- ton Mifflin Co., Boston, xxiii+284 pp. BUTTRICK, P. L.
1914. The probable origin of the forests of the Black Hills of South Dakota. For- est Quart., 12:223-227. Byers, G. W.
1961. The cranefly genus Dolichopeza in North America. Univ. Kansas Sci. Bull., 42:665-924.
Cahalaxe, V. H.
1948. The status of mammals in the U. S. National Park System, 1947. Jour. Mamm., 29:247-259.
1951. A program for restoring extirpated mammals in the National Park Sys- tem. Jour. Mamm., 32:207-210.
1954. Status of the black-footed ferret. Jour. Mamm., 35:418-424. Gary, M.
1902. Some general remarks upon the dis- tribution of life in northwest Nebraska. Proc. Nebraska Ornith. Union, 3:63- 75. 1917. Life-zone investigations in Wyoming. N. Amer. Fauna, 42:1-95. Chamhers, K.
1948. Lead woodrat killed by squirrel. Out- door Life, 102:8. Charles, G.
1961. Trapped . . . but not for long. South
Dakota Conserv. Dig., 28:4-6. 1964. World's only full mount of Audubon Bighorn? South Dakota Conserv. Dig., 31:23. Chxircher, C. S.
1959. The specific status of the New World red fox. Jour. Mann., 40:513-520. Clark, F. H.
1938. Inheritance and linkage relations of mutant characters in the deer mouse, Peromyscus maniculatus. Contrib. Lab. Vert. Genetics, Univ. Michigan, 7:1-11. Clemexts, F. E., and R. W. Chaney
1937. En\ironment and life in the great plains. Suppl. Publ. Carnegie Inst. Washington, 24:1-154. Coats, G. W.
1945. True weasel stories. South Dakota Conserv. Dig., 12:10.
CocKRLTM, E. L., and K. L. Fitch
1952. Geographic variation in red-backed mice (genus Clethrionomys) in the southern Rocky Mountain region. Univ. Kansas Publ., Mus. Nat. Hist., 5:281-292.
COUES, E.
1875. A critical review of the North Amer- ican Saccomyidae. Proc. Acad. Nat. Sci. Philadelphia, pp. 272-327.
1877. Fur-bearing animals: a monograph of North American Mustelidae. . . . Misc. Publ. U. S. Geol. and Geog. Sur\'. Terr., 8:xiv+l-348.
CouES, E., and J. A. Allex
1877. Monographs of North American Ro- dentia. Rept. U. S. Geol. Surv, Terr., ll:xii-|-l-1091.
CowAx, I. M.
1940. Distribution and variation in the na- tive sheep of North America. Amer, Midland Nat., 24:508-580.
TURNER: MAMMALS OF THE BLACK HILLS
169
CULLBERTSON, T. A.
1952. Journal of an expedition to the Mau- vaises Terres and the upper Missouri in 1850. Smithsonian Inst., Bull. Bur. Amer. Ethnol., 147:viii+l-145.
Curtis, J. T.
1959. The vegetation of Wisconsin. Univ. Wisconsin Press, Madison, iii+657 pp.
CusHixG, J- E., Jr.
1945. Quaternary rodents and lagomorphs of San Josecito Cave, Nuevo Leon, Mexico. Jour. Mamm., 26:182-185.
1965. Problems in the Quaternary phyto- geography of the Great Lakes region. Pp. 403-416, in The Quaternary of the United States (H. E. Wright, Jr., and D. G. Frey, eds.), Princeton Univ. Press, vii-|-922 pp. Dartox, N. H.
1906. Geology of the Big Horn Mountains. Prof. Paper U. S. Geo!. Surv., 51: 1-129.
1909. Geology and water resources in the northern portion of the Black Hills and adjoining regions in South Da- kota and Wyoming. Prof. Paper U. S. Geol. Surv., 65:1-105.
Dartox, N. H., and S. Paige
1925. Central Black Hills folio. South Da- kota. Geol. Folio, U. S. Geol. Surv., 219:1-34. Deevey, E. S., Jr.
1949. Biogeography of the Pleistocene (Part I: Europe and North America). Bull. Geol. Soc. Amer., 60:1315-1416.
Dice, L. R.
1939. An estimate of the populations of deer-mice in the Black Hills of South Dakota and Wyoming. Contrib. Lab. Vert. Genetics, Univ. Michigan, 10: 1-5.
1941. Variation of the deer-mouse (Pero- myscus maniculatus) on the Sand Hills of Nebraska and adjacent areas. Contrib. Lab. Vert. Genetics, Univ. Michigan, 15:1-19.
1942. Variation in the deer-mouse (Pero- inijscus maniculatus) of the Badlands and Black Hills of South Dakota and Wyoming. Contrib. Lab. Vert. Ge- netics, Univ. Michigan, 19:1-10.
1947. Effectiveness of selection by owls of deer-mice (Peromyscus 7naniculatus) which contrast in color with their background. Contrib. Lab. Vert. Ge- netics, Univ. Micliigan, 34:1-20.
DiLLOX, L. S.
1956. Wisconsin climates and life zones in North America. Science, 123:167-176.
Dodge, R. I.
1876. The Black Hills. James Miller, Publ., New York, 151 pp.
DORF, E.
1959. Climatic changes of the past and pres- ent. Contrib. Mus. Paleo., Univ. Michigan, 13:181-210. Elley, W. B.
1945. History of the ])eaver in the Black Hills. South Dakota Conserv. Dig., 12:5. Erdbrixk, D. p.
1953. A review of fossil and Recent bears of the Old World. . . . Deventer, Jan de Lange, xii-|-597 pp. EvAxs, D.
1966. The magnificent elk. South Dakota Conserv. Dig., 12:5.
Fexxeman, N. M.
1931. Physiography of western United States. McGraw-Hill Book Co., Inc., xiii-f 534 pp.
Ferxald, M. L.
1935. Critical plants of the upper Great Lakes Region of Ontario and Michi- gan. Rhodora, 37:197-341.
FiXDLEY, J.S.
1953. Pleistocene Soricidae from San Jose- cito Cave, Nuevo Leon, Mexico. Univ. Kansas Publ., Mus. Nat. Hist., 5:633- 639.
1955. Speciation of the wandering shrew. Univ. Kansas Publ., Mus. Nat. Hist., 9:1-68.
1956. Distribution of some South Dakotan mammals. Nat. Hist. Misc., Chicago Acad. Sci., 155:1-2.
FixDLEY, J. S., and R. H. Baker
1956. Dwarf shrew first-reported in South Dakota. Jour. Mamm., 37:543.
FixDLEY, J. S., and C. Joxes
1964. Seasonal distribution of the hoary bat. Jour. Mamm., 45: 461-470.
Flint, R. F.
1955. Pleistocene geology of eastern South Dakota, Prof. Paper, U. S. Geol. Surv., 262:1-173.
1957. Glacial and Pleistocene Geology. John Wiley and Sons, Inc., New York, xiii -j-553 pp.
Frey, D. G.
1965. Other invertebrates — an essay in bio- geography. Pp. 613-631, in The Qua- ternary of the United States (H. E. Wright, Jr., and D. G. Frey, eds.), Princeton Univ. Press, vli-f 922 pp.
Froilaxd, S. G.
1962. The genus Salix (willows) in the Black Hills of South Dakota. Tech. Bull. U. S. Dept. Agric, 1269:1-75.
Fryxell, F. M.
1926. A new high altitudinal limit for the American bison. Jour. Mamm., 7:102- 109.
170
xMISCELLAXEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Gabriel, K. R.
1964. A procedure for testing the homoge- neity of all sets of means in analysis of variance. Biometrics, 20:459-477.
Garretsox, M. S.
1938. The American bison. New York Zool. Soc, New York, xii-j-254 pp. Garst, W. E.
1954. Black-footed Ferret in South Dakota. Jour. Mamm., 35:594.
Gastler, F. G., a. L. Maxon, and W. T. McKean
1951. Composition of some plants eaten by deer in the Black Hills of South Da- kota. Jour. Wildlife Mgt., 15:352-357. GiER, H. T.
1967. Vertebrates of the Flint Hills. Trans. Kansas Acad. Sci., 70:51-59.
GiLLILAXD, S.
1968a. Mountain goat hunters enjoy the sport. South Dakota Conserv. Dig., 35:3-4.
1968b. Beaver's old buck teeth. South Da- kota Conserv. Dig., 35:5-7. Glass, B. P.
1947. Geographic variation in Perognathtis hispidus. Jour. Mamm., 228:174-179. Gleasox, H. a.
1923. The vegetational history of the Mid- dle West. Ann. Assoc. Amer. Geog., 12:39-85.
GOLDMAX, E. a.
1910. Revision of the woodrats of the genus Neotoina. N. Amer. Fauna, 31:1-124.
1944. Classification of wolves. Pp. 387-507, in The wolves of North America (S. P. Young and E. A. Goldman), Amer. Wildlife Inst., Washington, D. C., xx +636 pp.
1950. Raccoons of North and Middle Amer- ica. N. Amer. Fauna, 60:vi+l-153. Graves, H. I.
1899. Black Hills forest reserve. Ann. Rept. U. S. Geol. Surv. (for 1897-98). 19: 67-164.
Greex, M.
1961. Pathologic vertebrate fossils and re- cent specimens. Proc. South Dakota Acad. Sci., 40:142-148.
1962. An arthritic grizzly bear from South Dakota. Proc. South Dakota Acad. Sci., 41:37-40.
Greex, M., and J. A. Lillegravex
1965. Significance of Rangifer in the Her- rick Formation of South Dakota. Proc. South Dakota Acad. Sci., 44:48-51.
Cries, J. P., and E. L. Txjllis
1955. The geologic history of the Black Hills. Pp. 31-35, in North Dakota Geol. Soc. Guidebook, South Dakota Black Hills Field Conf., Bismarck, North Dakota.
Griffith, C.
1958. Twentieth century buffalo hunt: Cus- ter State Park relives history of plains
each year. South Dakota Conserv. Dig., 25:16-19. Grixxell, G. B.
1875. Zoological report. Pp. 79-84, in Re- port of a reconnaissance of the Black Hills of Dakota made in the summer of 1874 (W. Ludlow), Dept. Engineers, U. S. Army, 121 pp.
1904. The mountain sheep and its ranges. Pp. 270-348, in American big game and its haunts. Forest and Stream Publ. Co., New York, 497 pp.
1928. Mountain sheep. Jour. Mamm., 9: 1-10.
1929. Pronghorn antelope. Jour. Mamm.. 10:135-141.
GUILDAY, J. E.
1967. The climatic significance of the Hos- terman's Pit Local Fauna, Centre County, Pennsylvania. Amer. Antiq- uity, 32:231. GuiLDAY, J. E., and E. K. Adam
1967. Small mammal remains from Jaguar Ca\'e, Lemhi County, Idaho. Tebiwa, 10:26-36. GuiLDAY, J. E., P. S. Martix, and A. D. McGrady
1964. New Paris No. 4: A late Pleistocene cave deposit in Bedford County, Penn- svlvania. Bull. Nat. Spel. Soc, 26: 121-194.
GUNIER, W. J.
1971. Long-distance record for movement of a gray bat. Bat Research News, 12:5. Hafstex, U.
1964. A standard pollen diagram for the Southern High Plains, U.S.A., cover- ing the period back to early Wiscon- sin glaciation. Rept. VI Int. Quater- narv Congr. (Warsaw, 1961), 2:407- 420.
Hall, E. R.
1936. Mustelid mammals from the Pleisto- cene of North America with systematic notes on some Recent members of the genera Mtistela, Taxidea and Mephi- tis. Publ. Carnegie Inst. Washington, 473:41-119.
1946. Mammals of Nevada. Uniw California Press, Berkeley, xi-f 710 pp.
1951. American weasels. Univ. Kansas Publ., Mus. Nat. Hist, 4:1-466.
1965. Names of species of North American mammals north of Mexico. Misc. Publ. Mus. Nat. Hist., Univ. Kansas, 43: 1-16.
Hall, E. R., and E. L. Cockrltm
1953. A synopsis of the North American microtine rodents. Univ. Kansas Publ., Mus. Nat. Hist., 5:373-498. Hall, E. R., and K. R. Kelsox
1951. Comments in the taxonomy and geo- graphic distribution of some North
TURNER: MAMMALS OF THE BLACK HILLS
171
American rabbits. Univ. Kansas Publ., Mus. Nat. Hist., 5:49-58. 1959. The mammals of North America. Ron- ald Press, New York, 2 vols. (l:xxx+ 1-546+79, 2:viii+547-1083+79). Halliday, W. E. D., and A. W. A. Brown
1943. The distribution of some important forest trees in Canada. Ecology, 24: 353-373.
Haxdley, C. O., Jr.
1959. A re\ision of American bats of the genera Euderma and Flccotus. Free. U. S. Nat. Mus., 110:95-246, 27 figs.
Hansen, Richard M.
1960. Age and reproductive characteristics of mountain pocket gophers in Colo- rado. Jour. Mamm., 41:323-335.
Hanson, Walter O.
1950. The mountain goat in South Dakota.
Ph.D. dissertation, Univ. Michigan,
viii4-91 pp. Harmon, W. H.
1944. Notes on mountain goats in the Black Hills. Jour. Mamm., 25:149-151.
Harris, D.
1952a. Supplemental deer feeding in the Black Hills. South Dakota Conserv. Dig., 19:6-7, 16.
1952b. Pauline— The blind fawn. South Da- kota Conserv. Dig., 19:12.
1952c. Motorists take deer toll. South Dakota Conserv. Dig., 19:15.
19.53. Snow in the Hills. South Dakota Con- serv. Dig., 20:16. HARms, D., and S. E. Aldous
1946. Beaver management in the northern Black Hills of South Dakota. Jour. Wildlife Mgt., 10:348-353.
Hay, O. p.
1923. The Pleistocene of North America and its vertebrated animals of the states east of the Mississippi River and from the Canadian provinces east of longi- tude 95°. Publ. Carnegie Inst. Wash- ington, 322:1-499.
The Pleistocene of the middle region of North America and its vertebrated animals. Publ. Carnegie Inst. Wash- ington, 322A: 1-385.
F. V.
Zoological report. Pp. 75-79, in Ex- ploration in the Dacota Country in the year 1855 (G. K. Warren), Exec. Doc, U. S. Senate, 34th Cong. 76: 1-79.
1859. Mammals. Pp. 706-711, in Explora- tions in Nebraska in the years 1855- 1857 (G. K. Warren), Exec. Doc, House of Representatives, 35th Cong., 2:620-747.
1862. Mammals. Pp. 138-151, in On the geology and natural history of the Upper Missouri (F. V. Hayden),
1924.
Hayden 1856.
Trans. Amer. Pliil. Soc, 12 (n.s.):
1-218. Hayward, H. E.
1928. Studies of plants in the Black Hills of
South Dakota. Bot. Gaz., 85:353-412. Henderson, F. R., P. F. Springer, and R. Adrian
1969. The black-footed ferret in South Da- kota. Dept. Came, Fish and Parks, Pierre, South Dakota, 37 unnumbered pp.
Henshaw, H. W.
1910. Seed-eating mammals in relation to
forestry. Ann. Rept. Dept. Agric. (for
1910), p. 551. Hibbard, C. W.
1949. Pleistocene vertebrate paleontology in
North America. Bull. Geol. Soc.
Amer., 60:1417-1428. 1960. An interpretation of Pliocene and
Pleistocene climates in North America.
Ann. Rept. Michigan Acad. Sci.. 62:
5-30. 1963. A late Illinoian Fauna from Kansas
and its climatic significance. Papers
Michigan Acad. Sci., Arts, and Letters,
48:187-221.
1970. Pleistocene mammalian local faunas from the Great Plains and Central Lowland Provinces of the United States. Pp. 395-433, in Pleistocene and Recent environments of the Cen- tral Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp.
Hibbard, C. W., and F. J. Hinds
1960. A radiocarbon date for a woodland musk-ox in Michigan. Papers Michi- gan Acad. Sci., Arts, and Letters, 45: 103-111. Hibbard, E. A.
1957. Range and spread of the gray and fox squirrels in North Dakota. Jour. Mamm., 37:525-531. Hill, R. R.
1946. Palatability ratings of Black Hills plants for white-tailed deer. Jour. Wildlife Mgt., 10:47-54. Hill, R. R., and D. Harris
1943. Food preferences of Black Hills deer. Jour. Wildlife Mgt., 7:233-234.
Hillman, C. N.
1968. Field observations of black-footed fer- rets in South Dakota. Trans. Amer. Wildlife and Res. Conf., 33:433-443.
Hodge, W. T.
1960. Climates of the states: South Dakota. U. S. Weather Bureau, Waslrington, D. C, 16 pp.
HODGINS, R. A.
1956. Weather conditions hamper deer hunters. South Dakota Conserv. Dig., 24:14.
172
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Hoffman, W. J.
1877. List of mammals found in the \-icinity of Grand River, D. T. [South Dakota]'. Proc. Boston Soc. Nat. Hist., 19:94- 102.
Hoffmann, R. S., and J. K. Jones, Jr.
1970. Influence of the Late-glacial and Post-glacial e\'ents on the distribution of Recent mammals on the Northern Great Plains. Pp. 355-394, in Pleisto- cene and Recent environments of the Central and Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), LTni\'. Press Kansas, Lawrence, 433 pp.
Hoffmann, R. S., and D. L. Pattie
1968. A guide to Montana mammals. Univ. Montana Printing Serv., x4-133 pp.
Hoffmann, R. S., and R. D. Taber
1960. Notes on Sorex in the Northern Rocky
Mountain alpine zone. jour. Mamm.,
41:230-234. Hoffmann, R. S., P. L. Wright, and F. E. Newby
1969. The distribution of some mammals in Montana. I. Mammals other than bats. Jour. Mamm., 50:579-604.
Hooper, E. T.
1952. A systematic review of the harvest mice (genus Reiihrodoniomtjs) of Latin America. Misc. Publ. Mus. Zool., Univ. Michigan, 77:1-255. HoNEss, R. F., and N. M. Frost
1942. A Wyoming big-horn sheep study. Bull. Wyoming Game and Fish Dept., l:vi+l-126. Howell, A. H.
1906. Revision of the skunks of the genus Spilogale. N. Amer. Fauna, 26:1-55.
1915. Revision of the American marmots. N. Amer. Fauna, 37:1-80.
1918. Revision of the American flying squir- rels. N. Amer. Fauna, 44:1-64.
1929. Revision of the American chipmunks (genera Tamias and Etitamias). N. Amer. Fauna, 52:1-157.
1938. Revision of the North American ground squirrels, with a classification of the North American Sciuridae. N. Amer. Fauna, 56:1-256. Jackson, D.
1966. Custer's Gold: the United States Cav- alry Expedition of 1874. Yale Univ. Press, New Haven, vi+152 pp. Jackson, H. H. T.
1928. A taxonomic review of the American long-tailed shrews ( genera Sorex and Microsorex). N. Amer. Fauna, 51: vi-t- 1-238.
1944. Big-game resources of the United States, 1937-1942. Res. Rept. U. S. Dept. Interior, 8:11-1+56.
1951. Classification of the races of the coy- ote. Pp. 227-341, in The clever coy-
ote (S. P. Young and H. H. T. Jack- son), Stackpole Co., Harrisburg, and Wildlife Mgt. Inst., Washington, D. C, xv+411pp.
Jakway, G. E.
1958. Pleistocene Lagomorpha and Roden- tia from the San Josecito Cave, Neuvo Leon, Mexico. Trans. Kansas Acad. Sci., 61:313-327.
Jenney, W. p.
1868. Report on the mineral wealth, climate and rainfall, and natural resources of the Black Hills of Dakota. Exec. Doc, U. S. Senate, 44th Cong., 51:1-71.
Jones, J. K. Jr.
1953. Geographic distribution of the pocket mouse, Perognathiis fasciatus. Univ. Kansas Publ., Mus. Nat. Hist., 5:515- 526.
1964. Distribution and taxonomy of mam- mals of Nebraska. Univ. Kansas Publ., Mus. Nat. Hist., 16:1-356. Jones, J. K., Jr., and H. H. Genoways
1967a. A new subspecies of the fringe-tailed bat, Mijoiis thijsanodes, from the Black Hills of South Dakota and Wy- oming. Jour. Mamm., 48:231-235.
1967b. Annotated checklist of bats of South Dakota. Trans. Kansas Acad. Sci., 70:184-196. Jones, J. K., Jr., and F. R. Henderson
1963. Noteworthy records of foxes from South Dakota. Jour. Mamm., 44:283. Jones, J. K., Jr., and B. Mursaloglu
1961. Geographic \ariation in the harvest mouse, Rcithrodoiitoinys megalotis, on the central Great Plains and in ad- jacent regions. Univ. Kansas Publ., Mus. Nat. Hist., 14:9-27. Jones, J. K., Jr., and R. L. Packard
1958. Myotis keenii septentrionalis in South Dakota. Jour. Mamm., 39:150. Kapp, R. D.
1970. Pollen analysis of Pre- Wisconsin sedi- ments from the Great Plains. Pp. 143- 155, in Pleistocene and Recent en- vironments of the Central Great Plains (W. Dort, Jr., and J. K. Jones, Jr., ed. ), Univ. Press Kansas, Lawrence, 433 pp.
Kellogg, R. S.
1956. Wliat and where are the wliitetails. Pp. 31-55, in The deer of North America . . . (W. P. Taylor, ed. ), Stackpole Co., Harrisbmg, and Wild- life Mgt. Inst., Washington, D. C, 668 pp.
Kendeigh, S. C.
1954. History and evaluation of various con- cepts of plant and animal communi- ties in North America. Ecolog>% 35: 152-171.
1961. Animal Ecology. Prentice-Hall Press,
TURNER: MAMMALS OF THE BLACK HILLS
173
Englewood Cliffs, New Jersey, \i+468 pp. Kexnicott, R.
1863. Description of four new species of SpcttyiopJiihis, in the collections of the Smithsonian Institution. Proc. Acad. Nat. Sci. Philadelphia, 15:157- 158. King, J. A.
1951. The subspecific identity of the Black Hills flying squirrels (Glouconitis sa- hrinus). Jour. Mamm., 32:469-470.
1955. Social beha\'ior, social organization, and population dynamics in a black- tailed prairie dog town in the Black Hills of South Dakota. Contrib. Lab. \'ert. Biol., Univ. Michigan, 67:1-123.
KOFORD, C. B.
1958. Prairie dogs, white faces, and blue grama. Wildlife Monogr., 3:1-78. Khanz, J. J., and L. E. Petersex
1970. Preliminary in\estigations into the use of aspen by wildlife, 1968-69 South Dakota. South Dakota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-75-R-ll:l-6-f-S. Krxjtzsch, p. H.
1954. North American jumping mice (genus Zapiis). Univ. Kansas Publ., Mus. Nat. Hist., 7:349-472. Lamster, E. R.
1943. A deer hunt in the Black Hills. South Dakota Conserv. Dig., 10:1, 14. L.\yxe, J. N.
1954. The biology of the red squirrel, Ta- miasciiirus hudsonictis loquax (Bangs), in central New York. Ecol. Monogr., 24:227-267. Lee, O. B.
1962. Strange visitors. South Dakota Con- serv. Dig., 29:21.
1965. Part moose? South Dakota Conserv. Dig., 32:19. Lehmer, D. J.
1970. Climate and culture history in the Middle Missouri Valley. Pp. 117-129, in Pleistocene and Recent environ- ments of the Central Great Plains ( W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 43.3 pp. Lemke, R. W., W. M. Laird, M. J. Tiptox, and R. M. Lindvall
1965. Quaternary geology of the Northern Great Plains. Pp. 15-27, in The Qua- ternary of the United States (H. E. Wright, Jr., and D. G. Frey, eds.), Princeton Univ. Press, vii-t-922 pp.
Leonard, A. B., and J. Frye
1954. Ecological conditions accompanying loess deposition in the Great Plains region of the United States. Join-. Geo!., 62:399-404.
Leopold, A. S.
1956. Too many deer. South Dakota Con- serv. Dig., 23:2-5. LoxG, C. A.
1965. The mammals of Wyoming. Univ. Kansas Publ., Mus. Nat. Hist., 14: 493-758. Ludlow, W.
1875. Report of a reconnaissance of the Black Hills of Dakota made in the summer of 1874. Dept. Engineers, U. S. Army, 121 pp. Lyox, M. W., Jr., and W. H. O.sgood
1909. Catalogue of the type-specimens of mammals in the United States Na- tional Museum, including the Bio- logical Survey collection. Bull. U. S. Nat. Mus., 62:x+l-325. MacGixitie, H. D.
1959. Climate since the late Cretaceous. Pp. 61-79, in Zoogeography (C. L. Hubbs, ed. ), Publ. Amer. Assoc. Adv. Sci., 51 :x-l- 1-509.
Maxx, T.
1959. The "phantom" of Elk Mountain. South Dakota Conserv. Dig., 29:3-5.
Martix^ p. S.
1959. Pleistocene ecology and biogeography of North America. Pp. 375-420, in Zoogeography (C. L. Hubbs, ed. ), Publ. Amer. Assoc. Adv. Sci., 51:x-l- 1-509.
Martix, R. a.
1971a. Small mammals from barn owl pellets.
South Dakota Bird Notes, 23:4-10. 1971b. New records of the dwarf shrew from South Dakota. Jour. Mamm., 52:835- 836. Maxwell, M. H., and L. N. Brown
1968. Ecological distribution of rodents on the High Plains of eastern Wyoming. Southwestern Nat., 13:143-1.58.
McCuLLOUGH, D. R.
1969. The Tule elk, its history, behaxior, and ecology. Univ. California Publ. Zool., 88: vii+ 1-209.
McIxTOSH, A. J.
1931. A botanical survey of the Black Hills
of South Dakota. Black Hills Eng.,
19:159-276. Mead, R. A.
1968. Reproduction in western forms of the
spotted skunk (genus Spilogale) . Jour.
Mamm., 49:373-390.
Mexgel, R. M.
1970. The North American Central Plains as an isolating agent in bird speciation. Pp. 279-340, in Pleistocene and Re- cent environments of the Central Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp.
174
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
1971. A study of dog-coyote hybrids and implications concerning hybridization in Canis. Jour. Mamm., 52:316-336. Merriam, C. H.
1888. Description of a new species of mea- dow mouse from the Black Hills of Dakota. Amer. Nat., 22:934-935.
1889. Description of a new marmot from the Black Hills of Dakota. N. Amer. Fauna, 2:7-9.
1891. Description of a new Evotomijs from
the Black Hills of South Dakota. N.
Amer. Fauna, 5:119. 1899. Life zones and crop zones of the
United States. Bull. U. S. Dept.
Agric, 10:7-79. 1918. Review of the grizzly and big brown
bears of North America ( genus Ur-
sus) with description of a new genus,
Vettilarctos. N. Amer. Fauna, 41:1-
136. Merwin, J.
1971. Rocky Mountain mule deer. South
Dakota Conserv. Dig., 38:33. Miller, A. H.
1941. Speciation in the avian genus Junco.
Univ. California Publ. Zool., 44:173-
434. Miller, G. S., Jr.
1897. Revision of the North American bats
of the family Vespertilionidae. N.
Amer. Faima, 13: 1-140. Miller, G. S., Jr., and G. M. Allex
1928. The American bats of the genera
Myotis and Pizonyx. Bull. U. S. Nat.
Mus., 144:viii-f 1-218.
MORGAX, L. H.
1968. The American beaver and his works. J. B. Lippincott and Co., Philadelphia, 330 pp.
Morris, R. D.
1969. Competiti\'e exclusion between Mi- crotiis and Clcthrionoiuys in the aspen parkland of Saskatchewan. Jour. Mamm., .50:291-301.
Moulthrop, p. N.
1936. Myotis volans interior in South Da- kota. Jour. Mamm., 17:413-414. Murie, O. J.
1951. The elk of North America. Stackpole Co., Harrisburg, and Wildlife Mgt. Inst., Washington, D. C, 376 pp. Murray, K. F.
1957. Pleistocene climate and the fauna of Burnet Cave, New Mexico. Ecology, 38:129-132. Nachtigal, R.
1961. Department introduces new game va- rieties. South Dakota Conserv. Dig., 28:4-5. Nelsox, E. W.
1909. The rabbits of North America. N. Amer. Fauna, 29:1-314.
1925. Status of the pronghorn antelope, 1922-1924. Bull. U. S. Dept. Agric, 1346:1-64. Newtox, H., and W. P. Jenney
1880. Report on the geolog>' and resources of the Black Hills. U. S. Geog. and Geol. Surv., Rocky Mtn. Region, Washington, D. C, xiv+555 pp. Oesch, R. D.
1967. A preliminary investigation of a Pleis- tocene vertebrate fauna from Crank- shaft Pit, Jefferson Countv, Missouri. Bull. Nat. Spel. Soc, 29:163-185. O'Hara, C. C.
1929. Custer's Black Hills Expedition of 1874. Black Hills Eng., 17:221-286. Osgood, W. H.
1909. Revision of the mice of the American genus Peromyscus. N. Amer. Fauna, 28:1-285. Over, W. H.
1932. Flora of South Dakota. Univ. South Dakota, Vermillion, 161 pp. Over, W. H., and E. P. Churchill
1945. Mammals of South Dakota. Univ. South Dakota Mus. (processed litera- tiue), .59 pp. Packard, R. L.
1960. Speciation and evolution of the pygmy mice, genus Baiomys. Uni\'. Kansas Publ., Mus. Nat. Hist., 9:579-670.
Palmer, T. S.
1897. The jackrabbits of the United States. Biol. Surv. Bull., U. S. Dept. Agric, 8( revised): 1-88.
Parikh, G. C.
1970. Epizootic hemorrhagic deer disease study, 1968-69 South Dakota. South Dakota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-75-R-11: 1-19+i.
Parmalee, P. W.
1967. A recent cave bone deposit in south- western Illinois. Bull. Nat. Spel. Soc, 21:119-147.
Parmalee, P. W., and D. F. Hoffmeister 1957. Archaeozoological evidence of the spotted skunk in Illinois. Jour. Mamm., 38:261.
Pearsox, O. p., M. R. Koford, and A. K.
Pearsox
1952. Reproduction of the lump-nosed bat (Corynorhiniis rajinesquei) in Cali- fornia. Jour. Mamm., 33:273-320.
Peck, S.
1964. A faunal survey of Wind Cave. Pp. 35-43 in Exploration in Wind Cave. Nat. Spel. Soc. Wind Cave E.xpedi- tion 1959, 50 pp.
Pettixgill, O. S., Jr., and N. R. Whitxey, Jr.
1965. Birds of the Black Hills. Spec. Publ. Cornell Lab., l:vii-|-l-139.
TURNER: MAMMALS OF THE BLACK HILLS
175
PiHTLE, E. C, and J. M. Laytox
196L Epizootic hemorrhagic disease in wliite-tailed deer — characteristics of the South Dakota strain of \iriis. Amer. Jour. Vet. Res., 22:104-108.
POPOWSKI, B.
1952. Stulvtailed tabbies. South Dakota Conserv. Dig., 19:2-3, 6. Potter, L. D., and D. L. Green
1964. Ecology of ponderosa pine in western North Dakota. Ecology, 45:10-22. PoTZGER, T- E., and B. C. Tharpe
1947. Pollen profile from a Texas bog. Ecol- og>-, 28:247-280. Progtjlske, D. R., and D. C. Duerre
1964. Factors influencing spotlighting counts of deer. Jour. Wildlife Mgt., 28: 27-34.
PnuiTT, W. O., Jr.
1954. Aging in the masked shrew, Sorex cinereus cinereus Kerr. Jour. Mamm., 35:35-39.
Rayxolds. W. F.
1868. The report of Brexet Brigadier Gen- eral W. F. Raynolds .... Exec. Doc, U. S. Senate, 40th Cong., 77:1-174.
RiCHARDSOX, A. H.
1969a. Annual spring browse utilization srn- vey — 1969. South Dakota Dept. Game, Fish and Parks, Pittman-Rob- ertson Proj. W-95-R-3:l-8.
1969b. 100,000 deer. South Dakota Conserv. Dig., 36:24-26. RiCHARDSOX, A. H., and T. R. DeMarce
1967. Using and improving pellet group counts to determine deer numbers in the Black Hills 1966-67, South Da- kota. South Dakota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-95-R-1 and 2:1-10.
RiCHARDSOX", A. H., and P. C. Russell
1970. Using and improving pellet group counts to determine deer numl)ers and trend of deer numbers in the Black Hills, 1969 South Dakota. South Da- kota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-95-R-4: 1-10+2.
RiCHMOXD, G. M.
1965. Glaciation of the Rocky Mountains. Pp. 217-230, in The Quaternary of the United States (H. E. Wright, Jr., and D. G. Frey, eds.), Princeton Univ. Press, \ii+922 pp.
ROBERTSOX, P. B.
1971. Variation in populations of the mea- dow jumping mouse, Zapits hudsonius, on the Great Plains. M.A. thesis, Univ. Kansas, 28 pp.
Roe, F. G.
1951. The North American Buffalo .... Univ. Toronto Press, viii+957 pp.
Rose, B. J.
1970. An anahsis of the deer hunter report card returns, 1968 South Dakota. South Dakota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-95- R-3: 1-3+ J3. Ross, H. H.
1965. Pleistocene events and insects. Pp. 583-596, in The Quaternary of the United States (H. E. Wright, Jr., and D. G. Frey, eds.), Princeton Univ. Press, vii+922 pp.
1970. The ecological history of the Great Plains: e\'idence from grassland in- sects. Pp. 225-240, in Pleistocene and Recent en\ironments of the Central Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp. RUDD, V. E.
1951. Geographical affinities of the flora of North Dakota. Amer. Midland Nat., 45:722-739.
RUHE, R. V.
1970. Soils, paleosols, and environment. Pp. 37-52, in Pleistocene and Recent en- \'ironments of the Central Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp.
Rydberg, P. A.
1896. Flora of the Black Hills of South Da- kota. Contrib. U. S. Nat. Herbarium, 3:463-523.
SCHAXTZ, V. S.
1946. A new badger from South Dakota.
Proc. Biol. Soc. Washington, 59:81-82. 1953. Additional records of the spotted
skunk in South Dakota. Jour. Mamm.,
34:124-125.
Schara, R.
1967. Too few mountains, too many goats. South Dakota Conserv. Dig., 34:22-24.
SCHXEEWEIS, J. C.
1968. A food habits study of whitetail deer in the Northern Black Hills. M.S. thesis South Dakota State Univ., 59 pp.
SCHULTZ, C. B.
1934. The Pleistocene mammals of Nebraska. Pp. 357-392, in The Geology and mammalian fauna of the Pleistocene of Nebraska. Bull. Nebraska State Mus., 1:319-392.
SCHIJLTZ, C. B., G. C. LUENINGHOENER, End
W. D. Fraxkforter
1951. A graphic resume of the Pleistocene of Nebraska. Bull. Nebraska State Mus., 3:1-41.
ScHULTz, C. B., and H. T. U. Smith (eds.) 1965. Upper Mississippi Valley Guide-book. INQUA Field Conf. C, 7th Cong.,
176
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
Internat. Assoc. Quaternary Res., 126 pp. ScHWARz, E., and H. K. Schto^arz
1943. The wild and commensal stock of the house mouse, Mus musctilus Linnaeus. Jour. Mamm., 24:59-72.
Sears, P. B.
1961. A pollen profile from the grassland province. Science, 134:2038-2039. Semken, H. a., B. B. Miller, and J. B. Stevens
1964. Late Wisconsin woodland muskoxen in association with pollen and inverte- brates from Michigan. Jour. Paleo., 38:823-825.
Setox, E. T.
1929a. Cats, wolves, and foxes. Lives of game animals, Doubleday, Doran, and Co., Inc., New York, l:xxxix-1- 1-640.
1929b. Hoofed animals: deer, antelope, sheep, cattle, and peccary. Lives of game animals, Doubleday, Doran, and Co., Inc., New York, 3:xix+l-780.
1929c. Squirrels, rabbits, armadillo, and opos- sum. Lives of game animals. Double- day, Doran, and Co., Inc., New York, 4:.x.xii+l-949.
Setzer, H. W.
1949. Subspeciation in the kangaroo rat, Dipodomijs ordii. Univ. Kansas Publ., Mus. Nat. Hist., 23:473-573.
Shlmek, B.
1948. The plant geography of Iowa. LTniv. Iowa Studies Nat. Hist., 18:1-178.
Shope, R. E., L. G. MacNamara, and R. Max- gold
1960. A virus-induced epizootic hemor- rhagic disease of the Virginia white- tailed deer (OdocoiJeus virp.inianus) . Jour. Expt. Med., 111:155-170. Silver, J.
1924. Rodent enemies of fruit and shade trees. Jour. Mamm., 5:165-173.
1941. The house rat. Circ. U. S. Fish and Wildlife Serv., 6:iv-fl-17.
SiMPsox, G. G.
194-3. Mammals and the nature of conti- nents. Amer. Jour. Sci., 24:1-31. Skixxer, M. F.
1942. The faima of Papago Springs Cave, Arizona, and a study of Stockoceros. Bull. Amer. Mus. Nat. Hist., 80:143- 220.
Smith, P. W.
1957. An analysis of post- Wisconsin bio- geography of the Prairie Peninsula region based on distributional phe- nomena among terrestrial vertebrate populations. Ecology, 38:205-218.
196.5. Recent adjustments in animal ranges. Pp. 633-642, in The Quaternary of the
United States (H. E. Wright, Jr., and
D. G. Frey, eds. ) , Princeton Univ.
Press, vii+922 pp. Stearns, C. E.
1942. A fossil marmot from New Mexico and
its climatic significance. Amer. Jour.
Sci., 240:862-878. Stebler, a. M.
1939. An ecological study of the mammals
of the Badlands and the Black Hills of
South Dakota and Wyoming. Ecology,
20:382-393. Stewart, O. C.
1951. Burning and natural vegetation in the
United States. Geog. Rev., 41:317-
320. Stewart, R. E., and C. A. Thilenius
1964. Black Hills Lakes and Streams. Lake and Stream classification report. South Dakota Dept. Game, Fish and Parks, Dingell- Johnson Proj. F-l-R-13, Job Nos. 14 and 15:1-101.
SuTTOx, D. A., and C. F. Nadler
1969. Chromosomes of the North American chipmunk, genus Eutamias. Jour. Mamm., 50:524-535.
SWEXK, M. H.
1908. A preliminary review of the mammals of Nebraska, with synopses. Proc. Nebraska Acad. Sci., 8:61-144.
Swift, L. W.
1941. Rock}' Mountain Goats in the Black Hills of South Dakota. Trans. N. Amer. Wildlife Conf., 5:441-443.
Thomas, E. S.
1951. Distri])ution of Ohio animals. Ohio Jour. Sci., 51:153-167.
Thompsox', J. B.
1965. Correspondence. Bat Research News, 6:37.
Thompsox, L. F., and D. A. Hausle
1971. Using and improving pellet group counts to determine deer numbers and trend of deer numbers in the Black Hills, 1970 South Dakota. South Dakota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-95-R-5: 1-8+3.
Thwaites, R. G.
1906. Maximilian, Prince of Wied's travels in the interior of North America, 1832- 1834. The Arthur H. Clark Co., Cleve- land. Early Western Travels, 1748- 1846,22:1-393.
Tomaxek, G. W., and G. K. Hulett
1970. Effects of historical droughts on grass- land vegetation in the Central Great Plains. Pp. 201-210, in Pleistocene and Recent enxironments of the Cen- tral Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp.
TURNER: MAMMALS OK THE BLACK HILLS
177
Trainer, D. O.
1964. Epizootic hemorrhagic disease in deer. Jour. Wildlife Mgt., 28:377-381. Traxseau, E.N.
1935. The prairie peninsula. Ecology, 16: 423-437. Tryon, C. a., Jr.
1947. The biolog>- of the pocket gopher (Tlioinomys ialpoidcs) in Montana. Tech. Bull. Agric. Exp. Sta., Montana State Coll., 448:1-30. TuLLis, E. L.
1951. The igneous and metamorphic geology of the Black Hills. Pp. 84-85, in Guide Book Fiftli Field Cong., Soc. Vert. Paleo. in Western South Da- kota (J. D. Bump, ed.), Mus. Geol., South Dakota School of Mines and Tech., 87 pp. Turner, R. W., and W. H. Davis
1970. Bats from die Black Hills of South Dakota. Trans. Kansas Acad. Sci., 72:360-364. Turner, R. W., and J. K. Jones, Jr.
1968. Additional notes on bats from western South Dakota. Southwestern Nat., 13: 444-458.
Tyers, J.
1963. Correspondence. Bat Banding News, 4:17. Udvardy, M. D. F.
1969. Dynamic zoogeography, with special reference to land animals. Van Nos- strand Reinhold Co., xviii+445 pp.
Van Gelder, R. G.
1959. A taxonomic revision of the spotted skunks ( genus Sp//oga/e ) . Bull. Amer. Mus. Nat. Hist., 117:229-392. Visher, S. S.
1914. A preliminary report on the biology of Harding County, northwestern South Dakota. Bull. Soutli Dakota Geol. Surv., 6:1-126. Voss, J.
1939. Forests of the Yarmouth and Sanga- mon Interglacial period in Illinois. Ecolog>', 20:517-528.
^^'ADE, M.
1947. The Oregon Trail Journal 1846. Pp. 383-511, in The journals of Francis Parkman. Harper and Brothers Publ., New York, 718 pp.
Warren, G. K.
1856. Explorations in the Dacota Country
in the year 1855. Exec. Doc, U. S.
Senate, 34th Cong., 76:1-79. 1859. Explorations in Nebraska in the years
1855-1857. Exec. Doc, House of
Representatives, 35th Cong., 2:620-
747.
Watts, W. A.
1967. Late plant macrofossils from Minne- sota. Pp. 59-88, in Quaternary Paleo-
ecology (E. J. Gushing and H. E.
Wright, Jr., eds.), Yale Univ. Press,
vii+433 pp. Watts, W. A., and R. C. Bright
1968. Pollen, seed and mollusk analysis of a
sediment core from Pickerel Lake,
northeastern South Dakota. Bull.
Geol. Soc. Amer., 79:855-876. Watts, W. A., and H. E. Wright, Jr.
1966. Late- Wisconsin pollen and seed analy- sis from tlie Nebraska Sandhills. Ecol- ogy, 47:202-210.
Were, W. A.
1965. Plant geography in the southern Rocky Mountains. Pp. 453-468, in The Quaternary of the United States (H. E. Wright, Jr., and D. G. Frey, eds.), Princeton Univ. Press, vii+922 pp. Wells, P. V.
1965. Scarp woodlands, transported grass- land soils, and concepts of grassland climate in the Great Plains region. Science, 148:246-249.
1970a. Postglacial vegetational history of the Great Plains. Science, 167:1574-1582.
1970b. Vegetational history of the Great Plains: a post-glacial record of conif- erous woodland in southeastern Wy- oming. Pp. 185-202, in Pleistocene and Recent en\'ironments of the Cen- tral Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp.
1970c. Historical factors controlling vegeta- tion patterns and floristic distribu- tions in the Central Plains region of North America. Pp. 211-221, in Pleistocene and Recent en\ironments of the Central Great Plains ( W. Dort, Jr., and J. K. Jones, Jr., eds.), Univ. Press Kansas, Lawrence, 433 pp. West, D. R.
1970. Locker and field checks of antelope and analysis of hunter report card data, 1968 South Dakota. South Da- kota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-95-R-3:l- 5+8.
1971. Spring inventory of antelope in South Dakota, 1970 South Dakota. South Dakota Dept. Game, Fish and Parks, Pittman-Robertson Proj. W-95-R-4:l- 4+6.
Westin, F. C, L. F. Puhr, and G. J. Buntley
1967. Soils of South Dakota. Soil Surv. Ser., Agric. E.xp. Sta., South Dakota State Univ., 3:1-32.
Wetmore, cm.
1968. Lichens of the Black Hills of South Dakota and Wyoming. Publ. Mus., Micliigan State Univ., Biol. Ser., 3: 209-464.
178
MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY
White, E. M.
1961. Drainage alignment of western South
Dakota. Amer. Jour. Sci., 259:207-
210. White, J. A.
1952. A new cliipmunk (genus Eutatuias)
from the Black Hills. Univ. Kansas
Publ., Mus. Nat. Hist., 5:259-262. 1953a. Genera and subgenera of chipmunks.
Univ. Kansas Publ., Mus. Nat. Hist.,
5:543-561. 1953b. Geographic distribution and taxonomy
of the chipmunks of Wyoming. Univ.
Kansas Publ., Mus. Nat. Hist., 5:583-
610. Wilson, R. L.
1967. The Pleistocene vertebrates of Michi- gan. Papers Michigan Acad. Sci., Arts,
and Letters, 52:197-234.
WiNCHELL, N. H.
1875. Geological report. Pp. 21-66, in Re- port of a reconnaissance of the Black Hills of Dakota made in tlie summer of 1874 (W. Ludlow), Dept. Engi- neers, U. S. Army, 121 pp. Woodward, H. R.
1930. Abnormal growth of incisors in the Fiber zihethicus. Proc. South Dakota Acad. Sci., 13:108-111.
WOOSTER, L. D.
1935. Notes on the effects of drought on animal populations in western Kansas. Trans. Kansas Acad. Sci., 38:351-352.
1939.
Wright, 1968.
1970.
Young, S 1944
1946a.
1946b,
1958.
The effects of drouth on rodent popu- lations. Turtox News, 17:26-27.
H.E.,jR.
History of the Prairie Penninsula. Pp. 78-88, in The Quaternary of Illinois (R. E. Bergstrom, ed. ), Spec. Publ. Univ. Illinois Coll. Agric, 14:1-179. Vegetational liistory of the Central Plains. Pp. 157-172, in Pleistocene and Recent environments of the Cen- tral Great Plains (W. Dort, Jr., and J. K. Jones, Jr., eds. ), Univ. Press Kansas, Lawrence, 433 pp. .P.
Their history, life habits, economic status, and control. Pp. 1-386, in The Wolves of North America (S. P. Young and E. A. Goldman), Amer. Wildlife Inst., Washington, D. C, XX-}- 636 pp.
The wolf in North American history. The Caxton Printers, Ltd., Caldwell, Idaho, 149 pp.
History, life habits, economic status, and control. Pp. 1-173, in The puma, mysterious American cat (S. P. Young and E. A. Goldman), Amer. Wildhfe Inst., Washington, D. C, xiv-|-358 pp.
The bobcat of North America .... Stackpole Co., Harrisburg, and Wild- life Mgt. Inst., Washington, D. C, 193 pp.
AVAILABLE MISCELLANEOUS PUBLICATIONS IN MAMMALOGY UNIVERSITY OF KANSAS MUSEUM OF NATURAL HISTORY
7. Handbook of mammals of Kansas. By E. Raymond Hall. Pp. 1-303, illustrated. December 13, 1955. Paper bound $1.50 postpaid.
18. Furbearers in Kansas: A guide to trapping. By Howard J. Stains and Rollin H. Baker. Pp. 1-100, 2 plates, 13 figures in text. November 19, 1958. Paper boimd 50 cents postpaid.
30. Collecting and preparing study specimens of vertebrates. By E. Raymond Hall. Pp. 1-46, 34 figures in text. May 21, 1962. Paper bound 75 cents postpaid.
34. Habits of the red fox in northeastern Kansas. By WUham C. Stanley. Pp. 1-31,
3 plates, 3 figures in text. December 21, 1963. Paper bound 25 cents postpaid.
39. The status of deer in Kansas. By Donald D. Anderson. Pp. 1-36. September 28, 1964. Paper bound 25 cents postpaid.
43. Names of species of North American Mammals north of Mexico. By E. Raymond Hall. Pp. 1-16. December 10, 1965. Paper bound 25 cents postpaid.
46. The distributional status of bats in Kansas. By J. Knox Jones, Jr., Eugene D. Fleharty, and Patrick B. Dunnigan. Pp. 1-33, 11 figures in text. May 1, 1967. Paper bound 25 cents postpaid.
49. Natural history of the prairie dog in Kansas. By Ronald E. Smith. Pp. 1-39,
4 plates, 10 figures in text. September 27, 1967. Paper bound 50 cents postpaid.
51. Contributions in mammalogy. A volume honoring Professor E. Raymond Hall. Edited by J. Knox Jones, Jr. Pp. 1-428, 122 figs. July 11, 1969. Paper bound $5.00 postpaid.
54. The dentition of glossophagine bats: development, morphological characters, variation, pathology, and evolution. By Carlton J. Phillips. Pp. 1-138, 49 figs. September 24, 1971. Paper bound $5.00 postpaid.
56. Systematics of the chiropteran family Mormoopidae. By James Dale Smith. Pp. 1-132, 40 figs. March 10, 1972. Paper bound $5.00 postpaid.
58. Systematics of three species of woodrats (genus Neotoma) in Central North America. By Elmer C. Bimey. Pp. 1-173, 44 figs. April 13, 1973. Paper bound $7.50 postpaid.
60. Mammals of the Black Hills of South Dakota and Wyoming. By Ronald W. Turner. Pp. 1-178, 14 figs. April 3, 1974. Paper bound $7.50 postpaid.
MONOGRAPHS IN MAMMALOGY
2. Readings in mammalogy. By J. Knox Jones, Jr. and Sydney Anderson. Pp. ix + 586. 1970. Paper bound $8.00 postpaid.
3. The distribution of mammals in Colorado. By David M. Armstrong. Pp. x + 415, 8 pis. Cloth bound $16.00 postpaid.
1
I4a
dookbindlrtg Co.; InCj
3 2044 093 361 624
Date Due
i
f