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UMl

HARVARD UNIVERSITY

LIBRARY

OF THE

Museum of Comparative Zoology

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

Ronald W. Turner

UNIVERSITY OF KANSAS

LAWRENCE 1974 April 3, 1974

UNIVERSITY OF KANSAS PUBLICATIONS
MUSEUM OF NATURAL HISTORY

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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

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

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

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

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-

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

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

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

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).

MISCELLANEOUS PUBLICATION MUSEUM OF NATURAL HISTORY

I0 4°

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

104'

NEWCASTLE

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).

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

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).

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

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.

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

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).

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

60-

H- 40-

z
o

z
<

^ 20

'j^

-f

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-

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

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

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

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-

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

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-

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

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-

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

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.

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

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

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

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,

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

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

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

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-

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

c u

H o

o c

f«^

^ be

t? bO

"rt C

o c

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

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

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

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

±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

±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

±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

±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

±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

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

color

Percent

red

reflectance

Percent

green

reflectance

Percent

blue

reflectance

19 (14^,59)

Mean

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

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

17.7
±4.73

Sarpy County, Nebraska

56.3 24.6

±0.40 ±0.21

19.1
±0.22

9(35,69)

Mean

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

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

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

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

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

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

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-

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

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,

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

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

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

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-

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

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-

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

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

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

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-

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

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*

n3 CO

a
o

C
fuqn

PL, W) S

« 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

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

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.

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

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)

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

4-'

c «

Si y

c o

Oj cj

O 0^

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

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

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

±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

±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

±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

±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

±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

±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

111

C
03

a
o

PL, bc S!

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

Harding County, South Dakota

45.0 45.3 30.1 24.6

±6.89 ±0.22 ±0.11 ±0.15

3 (U,29)

Mean

Bennett County, South Dakota

32.8 47.2 28.4 24.4

±4.65 ±0.32 ±0.12 ±0.19

7(25,59)

Mean

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

Northern and western Nebraska (see text)
49.5 45.5 29.7 24.8

±6.09 ±0.17 ±0.11 ±0.07

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

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

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

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

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

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

7 (35,45)

Mean

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 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

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

i-C

of specimens

iS M

4~*

averaged

o c

o -^

S rt O

55 (24 5,31$ )

Mean

6 (25,4$)

Mean
SD

7 (3,5,45)

Mean
SD

27 (16 5,115)

Mean

4 (25,25 )

Mean

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

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-

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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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

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

color

Percent

red

reflectance

Percent

green

reflectance

Percent

blue

reflectance

26 (16,$, 10 9 )

Mean

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

26.2
±1.37

Crook County, Wyoming

48.4 28.3

±0.22 ±0.20

23.3
±0.12

5(45,19)

Mean

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

27.4
±2.39

Harding County, South Dakota

49.5 27.0

±0.32 ±0.16

23.5
±0.21

17 (75,109)

Mean

27.5
±3.25

Bennett County, South Dakota

49.5 26.4

±0.37 ±0.22

24.1
±0.19

22 (115, 119 )

Mean

30.1
±3.26

Cherry County, Nebraska

49.5 26.6

±0.26 ±0.18

23.9
±0.17

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

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

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

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

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

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

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

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.

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

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

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TURNER: MAMMALS OF THE BLACK HILLS

101

P M NEBRASCENaS

> P M NEBRASCENSIS

P M NEBRASCENSIS

P M LUTEUS

BLACK HILLS

HARDING CO,

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

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

±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

±5.15

±0.25 ±0.16 ±0.14

23 (13c?, 10 9 )

m. nehrascensis, Harding County,
South Dakota

Mean

30.4

50.7 25.3 24.0

±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

±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 >-,

ID
O

3
05

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

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

c B

o a;

C
O

ft 5b 2i

o a> 0;

OJ ,s

Ph^ ^

23 (85,15$)

N. c. orolestes. Black Hills, South

Dakota and Wyoming

Mean

27.2 48.5 26.3 25.2

±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

±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

±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

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

±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

±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

±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

*-i

Ifoot
th

itest

nil

)matie
dth

ral
th

bdoidal
dth

iS w^

-~ tc

c « ^ S^' a3 ^'^ a o ^ S^ s «

.— o

averaged

e2^

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

±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

±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

±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

0)
J5

C
O

S
O

"Sb
c
o

'c 3

c
o

•n

:a
&

5c
o

tpd.^p

lo CO

CD
CO CO

^ CO

d d

u"ns

^ +1

A\o.i-q}ooj

00 M

1^ CO

02
CD (M

X.IB[[IXUIU

d d

CD d

JO ijiSuaq

mSuar

t^ TP

00 CO

CD CO

00 d

[B-Hsoy

+ 1

1— (

ipptJOKj

^. ^

in in

cq Ti^

oi d

(>i d

c4 d

piOpC[lUB'^

^ +1

o in

i-H

ippea-iq

dWv

15.6

±2.0

(M 00

o m

OpV!UT08X2

.5^ +1

c -^ +1

S
o

4-t

I[n>is JO

rt CO 00

'rt

CD 05

i[43u3[

Qt^d

^-'

i> d

53 1^ o

}S9;Ba.io

c
a.)
o

oi II

CA)

•*-t

c/f

ipSu9[

Hill

5.8
1.48

:z,

.iBa

tp' ^

'^ 1-H

^ +1

-" II

i{j§ua[

CO 00
-(■ d

CO I>

^ d

CD cq

d ^'

jooj puiH

oi II

ipguaj

CO ^

Tf< OQ

(M O

II«X

d ^

t^ d

ai in

in +1

in II

C<I

iflSugj

a CT)

CD O

in CO

00 00

d 00

F^ox

^+1

I-H

.lOIOO 10

O 00

.-1 05

1> 1>;

'-^ ci

in oi

c4 <N

auojL

oi II

<N II

(M II

C C aj

c8 .5 '^

"^ ^ 2

<u a^ 53

a '^ ^

3 O

I-H

05
CM

?— I

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

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

±2.97

±0.20 ±0.18

±0.16

20(115,95)

Harding County, South Dakota

Mean

21.8

48.1 27.3

24.6

±3.00

±0.26 ±0.18

±0.15

8(45,45)

Bennett County, South Dakota

Mean

23.6

47.6 26.8

25.6

±4.72

±0.28 ±0.21

±0.13

4(25,25)

Campbell Countv, Wyoming

Mean

19.9

52.7 24.6

22.7

±1.84

±0.24 ±0.13

±0.20

2(15,15)

Weston County, Wvoming

Mean

25.5

46.1 28.5

25.4

±2.12

±0.03 ±0.10

±0.06

5(45,15)

Converse County, Wyoming

Mean

21.2

48.1 27.4

24.5

±2.41

±0.42 ±0.38

±0.08

16 (85,85)

Niobrara Countv, Wyoming

Mean

23.2

51.3 25.7

23.0

±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

±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

±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

±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

±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

±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

±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

±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

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

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

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

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,

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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. ) .

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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,

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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

n2,

umb
buc

umb
kids

Z o

Z O

•43

O N
(In -S

;>>

>. 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

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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.

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1939. Forests of the Yarmouth and Sanga-
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1965. Scarp woodlands, transported grass-
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Univ. Kansas Publ., Mus. Nat. Hist.,

5:543-561.
1953b. Geographic distribution and taxonomy

of the chipmunks of Wyoming. Univ.

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1930. Abnormal growth of incisors in the
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1935. Notes on the effects of drought on
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Trans. Kansas Acad. Sci., 38:351-352.

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.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
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Inst., Washington, D. C, xiv-|-358 pp.

The bobcat of North America ....
Stackpole Co., Harrisburg, and Wild-
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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.

I4a

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