MAMMALS OF THE GRAND STAIRCASE-ESCALANTE NATIONAL MONUMENT: A LITERATURE AND MUSEUM SURVEY
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Monographs of the Western North American Naturalist 1, © 2002, pp. 1–64
MAMMALS OF THE GRAND STAIRCASE–ESCALANTE
NATIONAL MONUMENT:
A LITERATURE AND MUSEUM SURVEY
Jerran T. Flinders1, Duke S. Rogers1,2, Jackee L. Webber-Alston1, and Harry A. Barber3
ABSTRACT.—This is the first treatment of the mammals of the Grand Staircase–Escalante National Monument
(GSENM). GSENM was established in 1996 as a 1.7-million-acre (680,000-ha) federal land reserve under the jurisdic-
tion of the Bureau of Land Management (BLM). To successfully manage this new monument, the BLM is presently
developing a management action plan. To provide information for the proper management of mammal species of the
area, we have reviewed background literature for each mammal potentially found within the Monument boundaries. We
propose that a core area, surrounded by a buffer matrix, be used in GSENM and surrounding public lands to preserve
ecological processes in their natural state. One hundred thirteen mammalian species are categorized as follows: 82 con-
temporary species, 21 species questionably present, 4 introduced species reportedly in the Monument, and 6 historical
species. Altogether, potentially 107 mammalian species exist there currently. Of 82 contemporary species, 11 are listed
in the Utah State Sensitive Species List, 1 in the Convention on the International Trade in Endangered Species of Wild
Flora and Fauna (CITES), and 1 in the World Conservation Union (IUCN) Red Book. Mammals listed under the 1973
Endangered Species Act (ESA) are noted when applicable, as well as State of Utah listings as endangered, threatened,
sensitive, or extirpated. For each mammalian species listed, we present in telegraphic style a life history account, sensi-
tivity status, and currently accepted nomenclature.
Key words: mammals, Grand Staircase–Escalante National Monument, Utah, Mammalia, Rodentia, Carnivora, Artio-
dactyla, Insectivora, Chiroptera.
INTRODUCTION ference. Such extraordinary flora potentially
sustains unique species of animals. The Canyons
As part of the Colorado Plateau, the Grand of Escalante are a maze of winding, connecting
Staircase–Escalante National Monument (here- canyons of the Escalante River and its tribu-
after referred to as GSENM or the Monument) taries. These riparian canyons are vital corri-
is a fascinating region, brimming with biodiver- dors for many animals and sanctuaries for
sity. Little, however, is known of the mammals relict, isolated plant communities (Visitor Infor-
of the area, their distributions, and their popu- mation 1998).
lation densities. The Grand Staircase–Esca- The GSENM spans biotic zones, from semi-
lante area was established in 1996 under the arid deserts to coniferous forests, with sparse,
Federal Antiquities Act as a 1.7-million-acre scattered water sources in between (Presiden-
(688,000-ha) national monument under jurisdic- tial Proclamation 1996). The distinctness of
tion of the Bureau of Land Management (BLM). these zones allows unequaled opportunity for
The Monument spans 3 distinct physiographic scientific and ecological study. Approximately
regions of south central Utah (Fig. 1). The 880,000 acres (352,000 ha) of Wilderness Study
Grand Staircase region is a series of multi-col- Areas exist to preserve the most remote areas
ored cliffs, starting from the Grand Canyon rim within the Monument until a management
and ascending about 5500 feet to the south- agenda is established.
west. The “stairs” consist of the Pink Cliffs, Few records exist on mammal collections
Gray Cliffs, White Cliffs, Vermillion Cliffs, and and sightings in GSENM. The earliest records
Chocolate Cliffs. The Kaiparowits Plateau is a of mammalian wildlife in the Monument are
remote area of mesas and steep cliffs. Because rock art (Fig. 2) left by ancient Native Ameri-
of its remoteness, many species of plants have cans: bighorn sheep, deer, bison, and elk (Raw-
evolved in isolation, away from human inter- ley 1986). Although the Monument is given
1Department of Integrative Biology, Brigham Young University, Provo, UT 84602.
2Monte L. Bean Life Science Museum, Brigham Young University, Provo, UT 84602.
3Bureau of Land Management, 318 North 1st East, Kanab, UT 84741.
12 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1 Fig. 1. Three distinct physiographic regions of the Monument: Grand Staircase, Kaiparowits Plateau, and Canyons of Escalante. Escalante’s name, the Dominguez-Escalante gated the Colorado River and bypassed the party of 1776 did not pass through the present border of the Monument en route to Arizona. boundaries of GSENM. Rather, the party by- The discovery of gold in the late 1880s south passed present-day borders in southern Kane of Glen Canyon brought people farther into County. In this area they record eating “hares, the area. Speculation about geologic wealth of rabbits, and wild sheep.” the region brought a later influx into the The first nonnative settlements in the region Kaiparowits area. Mineral exploration, such as were established by Mormon pioneers in the petroleum, began in the 1920s, with oil drilling early 1860s. Explorations by 2 of these pioneers, starting in 1921. The first significant drilling Jacob Hamblin and John D. Lee, aided in im- in the Monument was in the Upper Valley proving knowledge of the area. John D. Lee incline in 1948. In the late 1950s Glen Canyon established the now abandoned settlements of Dam required better road access and U.S. Paria in 1870, one at the mouth of the Paria Highway 89 was constructed. This provided River in 1871, and Adairville in 1873. These accessibility to the GSENM, inasmuch as the sites are now located within the Monument. highway skirts around or through the southern In 1871 and 1886, John Wesley Powell navi- and western borders (Murdock et al. 1974).
2002] MAMMALS OF GSENM 3
Fig. 2. Rock art found in GSENM has contributed to knowledge of the cultures of early Native Americans who inhab-
ited the region.
Presnall (1938) conducted a survey of the records of occurrence for the species are avail-
mammals in Zion National Park, Bryce Canyon able, these headings are not included.
National Park, and the Cedar Breaks areas
surrounding the Monument. Durrant (1952) PRESENT STATUS OF GSENM
surveyed throughout the state for mammalian
occurrence, but relied heavily on Tanner’s Three main controversial issues plague the
(1940) small mammal work for the Kaiparowits new GSENM: (1) the designation of Wilderness
Plateau and Paria River basin. Pritchett (1962) Study Areas within the Monument, (2) exist-
collected mammals throughout Kane County. ing coal mining rights within the boundaries
Raines (1976), Atwood and Pritchett (1974), of the Monument, and (3) continuance of graz-
and Murdock et al. (1974) studied the fauna of ing allotments within the Monument. The first
all the Kaiparowits region. However, their col- issue, wilderness designations, strongly influ-
lection sites were located primarily on the east ences the other 2 major issues and thus will be
side of the Colorado River, and only 3 were first in this discussion.
located in GSENM. Hayward, Killpack, Cof- Wilderness is herein defined (Wilderness Act
fery, and Pritchett (unpublished data) kept 1964) as any designated area “without perma-
mammalian population records in the area of nent improvements or human habitation” where
the Monument between 1952 and 1960 as well. the “imprint of man’s work is substantially
More recently, surveys have been conducted unnoticeable” in contrast to areas where “man
for chiropteran species by the BLM in the and his own works dominate the landscape.” It
summers of 1997 and 1998. In the following is also described as a place that “contains eco-
sections describing species in GSENM, records logical, geological, or other scientific, educa-
of occurrence for each species are listed as tional, scenic, and historical value.” We propose
Specimens Examined for museum records of that, to preserve biodiversity, wilderness areas
specimens, and Additional Records for occur- be designated based on the criteria discussed
rences cited in the above literature. Where no by Davidson et al. (1996). Davidson et al. (1996)4 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1
described how wilderness designations on BLM off gene flow and divide population corridors
land in Utah are based on ensuring long-term (Mader 1984).
population viability of native species, maintain- Among the preserve areas proposed in the
ing critical ecological and evolutionary pro- Redrock Wilderness Act are the Grand Stair-
cesses, and preserving the full range of com- case, Escalante Canyons, and Kaiparowits
munities, successional stages, and environmen- Plateau, much of which are in the borders of
tal gradients. Wilderness should be preserved GSENM (Comparison of Utah Wilderness
with the understanding that it is easier and Bills 1996). The Utah Wilderness Coalition
more cost-effective to protect species in intact and Southern Utah Wilderness Alliance also
and thus functioning ecosystems. include the White Cliffs as proposed wilder-
Designated wilderness areas may serve as ness along with the above-mentioned areas
core zones for metapopulations of large mam- (Stegner 1990).
mals, particularly carnivores, that maintain Wilderness designation honors pre-existing
some genetic flow through public land buffer mining and grazing claims on the land. Fear of
zones, connected and serving as a formal or loss of mineral exploration leaves some cau-
informal regional reserve network. It must be tious of promoting wilderness designations in
recognized that large carnivores have large the GSENM, as future exploration would not
home ranges and can be expected to disperse be allowed under the Wilderness Act (1964).
over long distances in appropriate habitats. For However, these deposits, if found upon explo-
example, annual, individual home ranges in ration, are likely too remote and difficult to
the Rocky Mountains are on the order of 150 reach, and thus undoubtedly are more costly
km2 for black bear, more than 400 km2 for than profitable to find and to extract. Goerold
mountain lion and wolverine, and nearly 900 (1990) reported that coal resources of Utah are
km2 for grizzly bear (Noss et al. 1996). Because abundant. Yet, the high cost of extraction cou-
of their requirements for space, position in the pled with inexpensive reserves from neighbor-
food web, and need for some management ing states may hamper local expansion in the
protection, large mammalian predators have Monument anyway. Goerold (1990) also stated
been considered indicators of the health or that most current coal mines contain enough
integrity of ecosystems (e.g., Eisenberg 1980, reserves for long periods of production at pres-
Noss 1995). Long-term managment plans for ent rates or at increased outputs. He indicated
GSENM should allow the Monument to serve that the coal industry contributed about 4% to
as a refugium for native wildlife and plants Utah’s economy and oil and gas less than 2%.
(Newmark 1985, 1995, Davidson et al. 1996). Grazing, by law, cannot be phased out once
Critical areas of wilderness designation are cen- an area is designated wilderness. Grazing allot-
ters for native and endemic species. Eighty-six ments also can be increased if the grazing does
percent of Utah’s indigenous plant species are not have adverse effects on ecosystems. Wilder-
found in arid and semiarid areas like GSENM ness can even benefit livestock operations by
(Davidson et al. 1996). Many parts of GSENM decreasing accessibility to stock and thus de-
are low-elevation areas of extensive native creasing theft and harassment. Livestock graz-
species diversity, and thus vital for preservation. ing, however, if improperly managed in wilder-
Preservation of corridors for wildlife move- ness areas, may lead to ecological alterations
ment is a major benefit of wilderness designa- such as introductions of exotic species, soil ero-
tions for mammalian species. If corridors are sion, competition with indigenous species, and
lost, this creates an island effect on the diver- deterioration of water quality (LeGate 1990).
sity of species and could result in a genetic Since no ground truth studies have yet been
bottleneck and ultimately a loss of biodiversity. completed in GSENM, this paper relied heav-
Although public lands are not true isolates, ily on literature and museum records to create
patterns of mammalian extinction exceed the a species inventory. Historical locations of
number of colonizations, and the rate of ex- specimen records are sometimes vague; there-
tinction is inversely related to size of the pro- fore, no accurate distributions could be con-
tected area, as is the case of true isolates (e.g., structed for mammals in GSENM. Scanty and
Newmark 1995). Wilderness designation serves obscure data led to noting mammals as intro-
to maintain areas as pristine as possible, thus duced, probable, or questionable species pres-
limiting roads and other such barriers that cut ent in GSENM. Table 1 lists mammals that2002] MAMMALS OF GSENM 5
have voucher specimens or are documented as although monogamy was more common in all
being in GSENM in the literature. These are 4 groups (Thomas et al. 1993).
mammals known to be in the Monument, ECOLOGY.—From about 500 A.D. and for 8
whereas other mammals listed herein are centuries following, Fremont people lived in
included based on distributions reviewed in Utah, W Colorado, and E Nevada. Dwelt along
the literature. The following species accounts Escalante River, its tributaries, and Kaiparo-
are organized in a modified telegraphic style wits Plateau. Lived to the north of contempo-
under the headings Historical, Contemporary, raries, the Virgin River Anasazi, and were in-
Questionable, and Introduced Species. Non- fluenced greatly by their culture. Shared traits
GSENM mammals are noted by common name. were pithouses, surface adobe houses, figurines,
Plant species are indicated only by common and black-on-gray painted pottery. Fremont
name, following designations in Welsh et al. were adapted to life style of mixed hunting,
(1987). Scientific names of avian species also gathering, and farming. Range likely constrained
are not included; the species are referred to to water availability. Moved between farming
by accepted American Ornithological Union locations of watered canyons and winter homes
checklist names. All other species are referred in uplands where hunted game. By 14th cen-
to by common name. Table 2 identifies cita- tury the people had disappeared (Waldman
tions, and their acronyms, for specimens 1985, Fahey et al. 1995, McFadden 1997). Early
examined. Anasazi utilized cliffs and canyon walls as
homes. Subsisted on opportunistic diet includ-
SPECIES OF HISTORICAL ing yucca, fruits, nuts, berries, mushrooms,
OCCURRENCE mule deer, antelope, leporids, fish, turkey and
other birds, and rodents. Cultivated corn, cot-
Homo sapiens Linneaus, 1758 ton, and Indian rice grass. Anasazi consisted of
Particularly Fremont, Virgin River Anasazi, 2 distinct cultures in GSENM, the Kayenta and
Kayenta Anasazi, and Southern Paiute the Virgin Anasazi. In Grand Staircase area,
Native American Cultures Virgin Anasazi relied heavily on agriculture.
Patterns of settlement reflected residential
Homo sapiens Linneaus, 1758. Syst. Nat., 10th ed., 1:20. mobility that allowed changes in various agri-
Type locality in Uppsala, Sweden (Wilson and Reeder
1993).
cultural environments. Next Anasazi group mi-
grated from Kayenta area and occupied Kai-
GENERAL CHARACTERISTICS.—Characteris- parowits Plateau region after or immediately
tics pertain to those described as early North before disappearance of Fremont. Because of
American cultures of the SW U.S. adaptation to conditions, group likely had mixed
DISTRIBUTION.—The Fremont group lived agricultural and hunter/gather program like
in the Escalante area from 500 A.D. to 1100 Fremont (McFadden 1996, 1997). Early South-
A.D. Two cultures of Anasazi existed in ern Paiute people immigrated in ca 1300 A.D.
GSENM, the Kayenta Anasazi and the Virgin into GSENM. Jackrabbits were hunted by chas-
River Anasazi. The Virgin River Anasazi (100 ing them into long, low nets made from twisted
B.C.–1200 A.D.) dwelt on the Grand Staircase fiber cordage. Paiutes were hunters, fishers,
portion of the Monument, and the Kayenta and gatherers who ate diverse foods. Paiutes
Anasazi (1050–1200 A.D.) lived in the Escalante adept in knowing healing properties of certain
area, particularly the Kaiparowits Plateau, plants. Built valley homes of willow sticks and
toward the end of the Fremont occupation. woven tule reeds in summer and lived in
The Southern Paiute group was established in foothills in winter. Tule reeds also used to craft
S Utah following the Fremont and Anasazi boats and duck decoys for hunting (e.g., Wald-
(1300 A.D.; D.A. McFadden personal commu- man 1985, Josephy 1994, Fahey et al. 1995,
nication 1998). McFadden personal communication 1998).
REPRODUCTION.—Usually single, altricial REMARKS.—Treatment only for native tribes
young born after 9-month gestation. Mating prior to European contact. Also evidence of
system variable with tribes. Early Southern human presence before cultures listed. Remains
Paiute had a polyandrous mating system when found in GSENM dating back to 6000 B.C.
they were especially prosperous or when near Big Water (McFadden personal commu-
women were scarce. Polygamy was accepted, nication 1998).6 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1
TABLE 1. List of mammals with records in GSENM.
INSECTIVORA RODENTIA
Northern Water Shrew (Sorex palustris) White-tailed Antelope Squirrel
CHIROPTERA (Ammospermophilus leucurus)
Pallid Bat (Antrozous pallidus) Golden-mantled Ground Squirrel
Townsend’s Big-eared Bat (Corynorhinus townsendii) (Spermophilus lateralis)
Big Brown Bat (Eptesicus fuscus) Rock Squirrel (Spermophilus variegatus)
Spotted Bat (Euderma maculatum) Cliff Chipmunk (Tamias dorsalis)
Allen’s Big-eared Bat (Idionycteris phyllotis) Least Chipmunk (Tamias minimus)
Silver-haired Bat (Lasionycteris noctivagans) Colorado Chipmunk (Tamias quadrivittatus)1
Hoary Bat (Lasiurus cinereus) Hopi Chipmunk (Tamias rufus)
California Myotis (Myotis californicus) Uintah Chipmunk (Tamias umbrinus)
Long-eared Myotis (Myotis evotis) Valley Pocket Gopher (Thomomys bottae)
Fringed Myotis (Myotis thysanodes) Southern Pocket Gopher (Thomomys umbrinus)
Long-legged Myotis (Myotis volans) Ord’s Kangaroo Rat (Dipodomys ordii)
Yuma Myotis (Myotis yumaensis) Northern Grasshopper Mouse (Onychomys
Western Pipestrel (Pipistrellus hesperus) leucogaster)
Mexican Free-tailed Bat (Tadarida brasiliensis) Long-tailed Pocket Mouse (Chaetodipus formosus)
CARNIVORA Little Pocket Mouse (Perognathus longimembris)
Coyote (Canis latrans) Great Basin Pocket Mouse (Perognathus parvus)
Gray Fox (Urocyon cinereoargenteus) Montane Vole (Microtus montanus)
Red Fox (Vulpes vulpes) Bushy-tailed Woodrat (Neotoma cinerea)
Bobcat (Lynx rufus) Sonoran Woodrat (Neotoma devia)
Mountain Lion (Puma concolor) Desert Woodrat (Neotoma lepida)
Striped Skunk (Mephitis mephitis) Brush Mouse (Peromyscus boylii)
Long-tailed Weasel (Mustela frenata) Canyon Mouse (Peromyscus crinitus)
Badger (Taxidea taxus) Deer Mouse (Peromyscus maniculatus)
American Black Bear (Ursus americanus) Pinyon Mouse (Peromyscus truei)
Western Harvest Mouse (Reithrodontomys megalotis)
ARTIODACTYLA
Pronghorn (Antilocapra americana) LAGOMORPHA
Mule Deer (Odocoileus hemionus) Black-tailed Jackrabbit (Lepus californicus)
Bighorn Sheep (Ovis canadensis) Desert Cottontail (Sylvilagus audubonii)
1Recorded as this species, but name likely since changed.
TABLE 2. Citations for specimens examined.
Acronym Citation and location
BLM Bureau of Land Management. 1998 Grand Staircase–Escalante Monument bat survey
summary. June–July. Unpublished data.
BYU Monte L. Bean Life Science Museum. Brigham Young University, Provo, Utah.
MVZ Museum of Vertebrate Zoology, University of California–Berkeley, Berkeley, California.
USNM National Museum of Natural History, Vertebrate Zoology Department of U.S. Fish
and Wildlife Service, Washington, D.C.
UU Utah Museum of Natural History, University of Utah, Salt Lake City, Utah.
Canis lupus Linnaeus, 1758 and brown to coal black. However, coloration
usually grizzled gray. Males slightly larger than
Gray Wolf females. Total length: 1300–1835 mm; length
Canis lupus Linnaeus, 1758. Syst. Nat., 10th ed., 1:39. Type of tail: 300–450 mm; length of hind foot: 250–
locality Sweden (Mech 1974). 275 mm; body mass: 18–80 kg (Mech 1974,
Clark and Stromberg 1987).
GENERAL CHARACTERISTICS.—Largest mem- DISTRIBUTION.—Formerly throughout all
ber of the canid family. Fur long and coloration of Northern Hemisphere in all habitats and
variable from pure white through mottled gray topography except extreme deserts and high2002] MAMMALS OF GSENM 7
mountain tops, now only disjunct populations. thick, colored medium brown to black. Head,
Reintroductions in progress in North America saddle, tail, and legs usually darker with black-
for select portions of W U.S., such as Yellow- ish facial mask and lighter upper body stripe.
stone National Park (Mech 1974, Wilson and Stripe may be creamy to yellowish, light brown,
Reeder 1993). Originally statewide distribution or reddish; extends from head and shoulders to
in Utah, except west desert region; no known base of tail. Light patches on throat and chest
occurrence in Utah at present (Durrant 1952). also common. Cream-colored animals with
REPRODUCTION.—Mated pairs likely breed brown feet also have been trapped, but thought
for life. Breeding January to April with typical to be rare. Males larger than females. Total
copulation lasting up to 30 minutes. Gestation length: 940–1070; length of tail: 218–260;
ca 63 days with 1–11 young. Sexually mature length of hind foot: 178–190; body mass: 6.6–
at 2 years of age, but usually breeds at 3 years 16.2 kg (Pasitschniak-Arts and Lariviere 1995).
(Fitzgerald et al. 1994). Breeding controlled DISTRIBUTION.—Historically extended down
by social hierarchy in each pack. the Rocky Mountains from Alaska to New Mex-
ECOLOGY.—Historically occupied most hab- ico and Arizona. Extirpated by the 20th century,
itats across Northern Hemisphere. Primary but now reestablishing populations in Montana
predators of deer, moose, wapiti, caribou, musk- and rare sightings in Colorado. Commonly
ox, mountain sheep, mountain goat, domestic thought never abundant in all other W U.S.
livestock, and beaver. Most of time spent in states (Pasitschniak-Arts and Lariviere 1995).
search of prey. Prey located by scent, tracking, No records since 1950 in Utah. Formerly found
and chance. Can survive up to 2 weeks without in high Wasatch, Uinta, and Boulder Mountains
prey. Consumes all of prey except largest bones (Durrant 1952).
and fur. Social animals, living in packs of a few REPRODUCTION.—Apparently polygynous
wolves up to 30, consisting of at least 1 pair of mating system. Breeds in May to August.
breeding adults, pups, and extra adults. Packs Monestrous with delayed implantation. Partu-
establish linear dominance hierarchies for both rition in spring with ca 2–3 young. Young
males and females. Dominant animals have first weaned 7–8 weeks. Females reproduce after 2
selection in feeding, breeding, and bedding years and exclusively raise young (Pasitschniak-
sites; often lead the pack when hunting or Arts and Lariviere 1995).
traveling. Species establishes home ranges of ECOLOGY.—Found in remote areas of mature
variable sizes, larger home range for larger forests such as Douglas-fir, alpine fir, and
packs. Has no natural predators except man. lodgepole pine. Opportunistic feeder relying
Three main methods of communication: (1) heavily on carrion, but will kill own food. Fol-
howling and similar vocalizations, (2) visual lows predators such as gray wolf or lynx to
displays of postures and positions, (3) scent scavenge remains. Feeds on caribou, moose,
marking (Mech 1974, Fitzgerald et al. 1994). lemmings, shrews, voles, snowshoe hare, mag-
STATUS.—Registered in Appendix II in pies, other birds and fish, beaver, lynx, ground
CITES, U.S.A. ESA lists it as Endangered in squirrels, and any available carcass or small
the USA (48 conterminous states, excluding mammal. Though scavenger, reported to have
Minnesota); and IUCN lists it as vulnerable
successfully attacked and killed big game such
(Wilson and Reeder 1993). In Utah, listed as
as caribou. Kills larger prey by biting on back
extirpated on the Sensitive Species List (Kim-
of neck, back, and withers. Wolverine urine
ball 1997).
odor reported to affect feeding areas of mule
Gulo gulo (Linnaeus, 1758) deer and snowshoe hare. Gray wolf, black bear,
brown bear, cougar, and golden eagle are
Wolverine
potential predators, though most frequently
[Mustela] gulo Linnaeus, 1758. Syst. Nat., 10th ed., 1:45. killed by man. Solitary, nocturnal, large home
Type locality Sweden, Lapland (Pasitschniak-Arts and
Lariviere 1995). ranges, and no territorial defense. Caches food
when excess amount is available. Scent marking
GENERAL CHARACTERISTICS.—Largest muste- important form of communication (Pasitschniak-
lid. Large head, broad forehead, short stout Arts and Lariviere 1995).
neck, relatively short legs, arched back, and STATUS.—Listed by IUCN as vulnerable, and
heavy musculature. Short, round, well-furred in Utah listed as a threatened species (Wilson
ears. Small, beady eyes. Bushy tail. Pelage and Reeder 1993, Kimball 1997).8 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1
Mustela nigripes (Audubon Ursus arctos Linnaeus, 1758
and Bachman, 1851) Grizzly Bear
Black-footed Ferret Ursus arctos Linnaeus, 1758. Syst. Nat., 10th ed., 1:47. Type
locality “Sweden” (Pasitschniak-Arts 1993).
Putorius nigripes Audubon and Bachman, 1851. Viviparous
Quadrupeds of North America, 2:297. Type locality Fort
GENERAL CHARACTERISTICS.—Massive head
Laramie, Goshen Co., Wyoming (Wilson and Reeder
1993). with dished face, rounded ears, small eyes,
and prominent nose. Exhibits heavy shoulder
GENERAL CHARACTERISTICS.—Mink size and hump, short tail, and a powerful, large body.
shape. Upperparts yellowish buff or whitish. Coloration of pelage variable from tan, blond,
Venter paler. Feet, mask over eyes, and crown gold, gray, silver, cinnamon, and all shades of
of head black. Tail long and tipped with black. brown to almost black. Interior animals tend
Distinguished from weasels by blending of to be lighter than coastal and have pale-tipped
color changes instead of sharp boundaries. guard hairs, giving a grizzled appearance.
Females smaller than males. Total length: Head and shoulders usually paler than darker
500–533; length of tail: 114–127; length of hind sides, belly, and legs. Males larger than females
foot: 60–73; length of ear: 29–31; body mass: (Pasitschniak-Arts 1993). Total length: 1700–
2800; length of tail: 70–80; length of hind foot:
530–1300 g (Hillman and Clark 1980, Fitzgerald
230–280; length of ear: 100–130; body mass:
et al. 1994).
135–275 kg (Fitzgerald et al. 1994).
DISTRIBUTION.—Formerly throughout Great DISTRIBUTION.—Historically in W and C
Plains, mountain basins, and semiarid grass- North America from Arctic Ocean to C Mexico.
lands of North America, coinciding with prairie Now found in Eurasia, Alaska, Canada (except
dog range. Few remnant populations still mark open prairies), and disjunct populations in
the range (Hillman and Clark 1980). However, Washington, Montana, Idaho, and Wyoming.
viable populations only in captivity and cur- Extirpated in Mexico in 1960s, but possibly a
rent reintroduction efforts are underway on few in N Mexico (Pasitschniak-Arts 1993). Prior
the Colorado-Utah border (Wilson and Reeder to extinction in Utah, occurrence in Sanpete
1993, Bates personal communication 1999). and Washington counties as well as NE Utah
REPRODUCTION.—Breeding late February and SE Idaho borders (Durrant 1952, Russell
to early April. Gestation 42–45 days with 2–4 1955).
young born in May. Presumed polygynous sys- REPRODUCTION.—Polygamous system with
tem with sexual maturity reached at 1 year several males competing for estrous females.
(Hillman and Clark 1980, Fitzgerald et al. 1994). Breeding mid-May to July. Delayed implanta-
ECOLOGY.—In close association with occur- tion. Young born January to March in female
rence of prairie dog. Preys on prairie dogs and dens and stay with mother for 2–3 years.
uses their burrows for dens and shelter. Also Females bear young at ca 7 years (Pasitsch-
takes ground squirrels, cottontail rabbits, deer niak-Arts 1993).
mice, and small birds. Ferret number declines ECOLOGY.—Inhabits any area with sufficient
directly related to habitat destruction, sec- resource availability from prairie grassland to
alpine tundra. Today, however, found in more
ondary poisonings, and prairie dog population
remote areas where contact with people is lim-
control. Predators reported as coyote, golden
ited. Diet mainly vegetation such as grasses,
eagle, and great horned owl. Highly antagonis- succulent herbs, tender shoots, flowers, leaves,
tic to same sex conspecifics. Habits secretive roots, bulbs, tubers, mosses, horsetails, willows,
and nocturnal. Less active in winter and solitary and berries. Also takes insects, larvae, birds,
except when breeding in spring. Vocalizations eggs, acorns, cones, nuts, fish, small mammals,
include chatters, hisses, and whimpers. Odor moose, caribou, elk, deer, pronghorn, bison, big-
recognition aids in night travel (Hillman and horn sheep, mountain goat, and domestic live-
Clark 1980). stock. Caches food. Strictly terrestrial mam-
STATUS.—Listed in CITES (Appendix I), mal; does not climb trees like the black bear.
and Endangered in U.S.A. ESA (Wilson and Solitary except mother with young. Active in
Reeder 1993). Listed in the State of Utah as an day, but mainly at night. Dens in winter months
endangered species (Kimball 1997). in response to food scarcity. Establishes home2002] MAMMALS OF GSENM 9
ranges that overlap with conspecifics but shows ECOLOGY.—Estimates indicate 30–50 mil-
no territorial defense, but rather maintains lion bison occurred in North America prior to
range by mutual avoidance. Only natural re- European colonization but were nearly exter-
ported predator is Siberian tiger. Habitat loss, minated due to overhunting (Nowak 1991). In
genetic bottleneck due to isolation, habituation 1887, Hornaday reported just 541 bison re-
to humans, and illegal hunting are major human- mained from the vast herd (Shaw and Meagher
induced threats to populations. Low reproduc- 2000). Prehistoric distribution primarily central
tive rate and slow compensatory response to grasslands and northern parklands of North
these threats are serious problems. Intolerance America. Now reintroduced herds in habitats
and aggression in males during mating season ranging from semidesert to boreal forests where
is due to a nonfamilial bond experienced as suitable grazing is available. Foraging gener-
cubs. Communicates by visual, auditory, and ally nonselective grazing on grasses, sedges,
vocalizations, but relies heavily on olfaction and forbs. Many herds managed as domestic
(Pasitschniak-Arts 1993, Fitzgerald et al. 1994). livestock. Free-ranging wild herds now in
STATUS.—Listed in CITES as threatened in Alaska, Canada, Yellowstone National Park, and
USA (Wilson and Reeder 1993). Listed in the the Henry Mountains of southern Utah (Shaw
State of Utah as an extirpated species (Kimball and Meagher 2000). Wallows in dry or muddy
1997). soil. Predators man and gray wolf (Meagher
1986, Clark and Stromberg 1987).
Bison bison (Linnaeus, 1758) REMARKS.—Prehistoric rock art has been
American Bison found depicting bison at Johnson Canyon
Bos bison Linnaeus, 1758. Syst. Nat., 10th ed., 1:72. Type within GSENM in C Kane Co. (Rawley 1986).
locality “Mexico” (=C Kansas, “Quivira”), redesignated Presnall (1938) refers also to a herd in House
as Canadian River Valley, E New Mexico (Wilson and Rock Valley within GSENM in S Kane Co.,
Reeder 1993). brought in from Texas in 1905.
STATUS.—Bison b. athabascae listed in
GENERAL CHARACTERISTICS.—Largest North
CITES (Appendix I), in U.S.A. ESA as Endan-
American artiodactyl. Body massive, accentu-
gered (Wilson and Reeder 1993). Geist (1996)
ated by shoulder hump. Pelage brown, long
considers the species Bison bison to be with-
over forehead, neck, hump, and front quarters;
out subspecies; identified variations attributed
shorter over rump, tail, and hind legs. Tail nar-
to differences in nutrition.
row with tufted tip. Head large and neck thick
and short. Males and females possess horns that Cynomys parvidens (Allen, 1905)
curve upward and inward, tapering to sharp Utah Prairie Dog
tip. Eyes anterolateral. Legs short and stout.
Hooves black and circular shaped on bottom. Cynomys parvidens Allen, 1905. Bull. Mus. Sci., Brooklyn
Inst. Arts and Sci., 1:117–122. Type locality Buckskin
Females slightly smaller than males. Total Valley, Iron Co., Utah (Pizzimenti and Collier 1975).
length: 2130–3800; length of tail: 300–910;
length of hind foot: 500–680; length of ear: GENERAL CHARACTERISTICS.—Small prairie
110–150; body mass: 410–910 kg (Meagher dog with each hair composed of several colors.
1986, Clark and Stromberg 1987). Upperparts cinnamon-clay to buff and darker
DISTRIBUTION.—Formerly throughout North on rump. Underparts paler cinnamon or buff.
America from Canada and W Canada to N Whitish mouth, chin, and tail tip; diagnostic
Mexico. Reintroductions occurring within his- dark cheek patches. Total length: 305–360;
toric range (Meagher 1986). Fairly extensive length of tail: 30–60; length of hind foot: 55–
ranching of domesticated bison in U.S. and 66; length of ear: 12–16; body mass 650–1050
Canada. Found in Utah on Antelope Island g (Pizzimenti and Collier 1975, Jacquart 1986).
(Great Salt Lake) and in E Utah from the San DISTRIBUTION.—SC Utah (Durrant 1952,
Rafael Swell to the Henry Mountains (Durrant Wilson and Reeder 1993).
1952). REPRODUCTION.—Breeding early spring with
REPRODUCTION.—Breeding season late June– ca 4–5 young. Reproduction may be delayed
September, but strong seasonal variance. Ges- 2–4 weeks at higher elevations. Annual partu-
tation ca 285 days with usually 1 young born rition (Pizzimenti and Collier 1975).
in mid-April to May. Calves more ruddy pelage ECOLOGY.—Once widely distributed
(Meagher 1986). throughout range, now occurs only sparsely10 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1
along the Sevier River and E Iron Co. Because Sorex merriami Dobson, 1890
of poisoning and disease, populations have Merriam’s Shrew
dwindled from 95,000 in 1920 to 3500 in 1976.
Sorex merriami Dobson, 1890. Monogr. Insectivora, pt. 3
Dense vegetation northward and possible com- (Soricidae), fasc. 1, pl. 23. Type locality [1 1/2 mi] above
petition with Uinta ground squirrel are proba- Fort Custer [Hardin], Big Horn Co., Montana (Wilson
ble reasons for lack of successful dispersal and and Reeder 1993).
range expansions (Zeveloff 1988). Primary diet
mostly forbs and grasses. Habits are diurnal GENERAL CHARACTERISTICS.—Small shrew.
and colonial. Burrows similar to C. leucurus Pelage pale gray above, with paler flanks;
with little design of mound (Pizzimenti and whitish below. Long, distinctly bicolored tail
Collier 1975). (Jameson and Peeters 1988). Total length: 88–
STATUS.—Listed in U.S.A. ESA as Threat- 107; length of tail: 33–42; length of hind foot:
ened; IUCN as vulnerable; in Utah it is a 11–13; length of ear: 8–9; body mass: 4.4–6.5 g
threatened species (Wilson and Reeder 1993, (Armstrong and Jones 1971).
Kimball 1997). DISTRIBUTION.—All W United States; W
Oregon to S New Mexico (Jung and Hoffmann
SPECIES OF CONTEMPORARY 1981). Possibly statewide distribution in Utah
OCCURRENCE within sagebrush, mountain mahogany, and arid
grassland communities, often in association
INSECTIVORA with sagebrush vole (Sparks 1974, Zeveloff
Notiosorex crawfordi (Coues, 1877) 1988).
REPRODUCTION.—Pregnant females mid-
Desert Shrew March to early July. Males with enlarged testes
Notisorex Coues, 1877. Bull. U.S. Geol. Geogr. Surv. Terr., and prominent flank glands (reproductive
3:631. Type locality 2 mi above El Paso, El Paso Co., attractant) caught in March–June (Armstrong
Texas (Wilson and Reeder 1993).
and Jones 1971).
GENERAL CHARACTERISTICS.—Color grayish ECOLOGY.—Found in drier habitats, usually
with brown above. Short grayish tail, paler sagebrush, some grasslands, mountain mahog-
below. Three unicuspid teeth in upper jaw, any, pinyon, and pine/fir/aspen areas. Utilizes
unlike members of Sorex genus. Ears rela- microtine runways and burrows. Feeds on spi-
tively large. Prominent flank glands. Largest ders, beetles, larval lepidopterans, and ichneu-
shrew in North America. Total length: 81–90; monid wasps. Caterpillars most often used in
length of tail: 24–26; length of hind foot: 9–11; summer (Armstrong and Jones 1971).
length of ear: 8–9; body mass 3–5 g (Jameson Sorex monticolus Merriam, 1890
and Peeters 1988).
Montane Shrew
DISTRIBUTION.—Broad distribution through-
out SW U.S., C and N Mexico (Hall 1981). At Sorex monticolus Merriam, 1890. N. Amer. Fauna, 3:43,
September 11. Type locality San Francisco Mtn.,
the same latitude as this portion of Utah, there
11,500 ft. [3450 m], May 15, Coconino Co., Arizona
is a record for Nye Co., Nevada (Durrant 1952). (Hall 1981).
REPRODUCTION.—Breeds in warmer months.
Gestation length not known. Litter size 3–5 GENERAL CHARACTERISTICS.—Color varies
young (Armstrong and Jones 1972). with seasons. Summer: short fur, rust brown,
ECOLOGY.—Semidesert scrub association of paler below; winter: gray fur longer and darker
mesquite, agave, and scrub oaks. Does not re- above than below. Tail bicolored. Total length:
quire permanent water sources and uses a 111–120; length of tail: 46–55; length of hind
wide variety of food including most insects foot: 13–15; length of ear: 6–7; body mass: 4–7
and even carrion. Habitat cover variable from g (Jameson and Peeters 1988).
brush piles to woodrat dens. Predators mostly DISTRIBUTION.—Mesic habitats from Alaska
owls, especially great horned and barn owls throughout Rocky Mountains to New Mexico
(Armstrong and Jones 1972). with isolated populations in C Mexico (Fitz-
STATUS.—Utah State Sensitive Species List gerald et al. 1994). Found throughout Utah in
due to limited distribution (restricted to south- all mountain ranges and adjacent valleys (Dur-
ernmost portion of Utah; Kimball 1997). rant 1952).2002] MAMMALS OF GSENM 11 REPRODUCTION.—Breeds from April to length of hind foot: 18–22; body mass: 8–14 g August. More than 1 litter per year. Gestation (Beneski and Stinson 1987). 20–22 days with ca 6 young per litter (Fitzger- DISTRIBUTION.—Boreomontane distribution ald et al. 1994). throughout Rocky Mountain range and much ECOLOGY.—Found in mesic habitats of aspen, of Canada and Hudsonian life zones (Beneski willow, moist meadows of subalpine forests, and Stinson 1987). Found throughout Utah in and riparian communities. Greatly affected by permanent water source (Sparks 1974). temporal variation in groundcover; when her- REPRODUCTION.—Two to 3 young per litter baceous cover declines, increases use in adja- with a gestation
12 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1
CHIROPTERA nose. Total length: 89–112; length of tail: 35–
54; length of hind foot: 10–13; length of ear:
Antrozous pallidus (Le Conte, 1856)
30–41; length of forearm: 39–47; body mass:
Pallid Bat 7–12 g (Jameson and Peeters 1988).
Antrozous pallidus Le Conte, 1856. Proc. Acad. Nat. Sci. DISTRIBUTION.—W U.S. from Washington
Philadelphia, 14:248 (Wilson and Reeder 1993). to Idaho, Wyoming, Colorado, New Mexico,
Oklahoma, and Texas; scattered populations in
GENERAL CHARACTERISTICS.—Large bat. Missouri, Arkansas, Kentucky, West Virginia, and
Coloring pale beige above, nearly white Virginia (Jameson and Peeters 1988). Found in
below. Big ears separated at base. Total length: S Utah (Hasenyager 1980, Mollhagen and Bogan
90–113; length of tail: 40–47; length of hind 1997).
foot: 10–12; length of ear: 28–32; length of REPRODUCTION.—Copulation throughout
forearm: 48–60; body mass: 14–17 g (Fitzger-
winter in hibernacula. Females store sperm
ald et al. 1994).
until after hibernation, when fertilization occurs.
DISTRIBUTION.—Generally common through-
Single young born in early summer (Zeveloff
out riparian habitats in arid deserts and grass-
1988). Gestation ranges from 56 to 100 days,
lands of SW U.S. (Hermanson and O’Shea
1983). Distributed in E and S Utah (Sparks depending on climatic factors and tempera-
1974). ture variations (Schmidly 1991).
REPRODUCTION.—Breeding October to De- ECOLOGY.—Common in highlands of the
cember, with sperm stored in female repro- W U.S. in scrublands as well as pine, pinyon-
ductive tract until spring. Parturition May–June, juniper, and deciduous forests; uncommon in
female hanging upright while altricial young deserts. Feeds mostly on moths although will
are born breech. Weaned 6–8 weeks (Herman- take other insects in late evening. Roosts soli-
son and O’Shea 1983). tarily or in small groups (Zeveloff 1988). Species
ECOLOGY.—Characteristic of desert areas, very intolerant to disturbance. Shelter a limit-
less abundant in evergreens and mixed forests. ing factor in distribution (Hasenyager 1980).
Co-roosts at night with other species of bats. REMARKS.—Wilson and Reeder (1993) cite
Forms large maternity colonies. Preys on flight- no evidence on why Corynorhinus should not
less arthropods, moths, and sometimes small be used. Herein, Corynorhinus is chosen based
lizards and smaller bats. Predators include on parsimony analysis of the phylogeny of Ple-
snakes and owls. Echolocation main source of cotus by Tumlison and Douglas (1992).
orientation. Isolation calls show individual dis- ADDITIONAL RECORDS.—Kane Co.: Pink
tinctiveness and aid in localization of off- Cove, 1 (BLM); Pete’s Cove, 2 (BLM).
spring. Squabble notes and irritation buzzes STATUS.—Registered as sensitive species in
audible at night roosts (Hermanson and O’Shea Utah due to limited distribution and declining
1983). populations (Kimball 1997).
SPECIMENS EXAMINED.—Kane Co.: 42 mi S,
18 mi W, Paria River, 1 (USNM); mouth of Eptesicus fuscus
Paria Canyon, 1 (USNM); Paria, 1 (BYU). (de Beauvois, 1796)
ADDITIONAL RECORDS.—Kane Co: Paria Big Brown Bat
(Pritchett 1962); Paria, 2 (Hasenyager 1980); Vespertilio fuscus Palisot de Beauvois, 1796. Cat. Raisonne
Fin Little Spring, 3 (BLM); Pink Cove, 1 Mus. Peale Philadelphia, p. 18. Type locality “Les envi-
(BLM). rons de Philadelphia,” Pennsylvania (Kurta and Baker
1990).
Corynorhinus townsendii
(Cooper, 1837) GENERAL CHARACTERISTICS.—Large bat.
Townsend’s Big-eared Bat Pelage brown above, varying from light (in
deserts) to dark (in forests), usually glossy;
Plecotus townsendii Cooper, 1837. Ann. Lyc. Nat. Hist. belly paler, with hairs dark at base. Black wings
New York, 4:73, November, paratypes from Columbia
River, Oregon (Hall 1981). and membrane (not furred). Total length:
98–138; length of tail: 34–56; length of hind
GENERAL CHARACTERISTICS.—Pelage paler foot: 11–13; length of ear: not given; length of
gray or brown above, buff below. Largest eared forearm: 39–54; body mass: 12–20 g (Fitzger-
bat in U.S. Two distinct, glandular lumps on ald et al. 1994).2002] MAMMALS OF GSENM 13
DISTRIBUTION.—From S Canada to N Vene- ECOLOGY.—Wide variety of habitats but
zuela, Colombia, and Brazil. Found in arid W most often collected in dry desert terrains.
and SW U.S. and forested highlands (Kurta Commonly feeds on moths, but also June bee-
and Baker 1990). Records throughout whole tles, flies, mealworms, katydids, and grasshop-
state of Utah, with great variation in color pers. With moths, spotted bat generally pulls
(Hasenyager 1980). wings and head off before eating. Predators
REPRODUCTION.—Copulation occurs in Sep- most often include kestrel, red-tailed hawk,
tember to March, with ovulation and fertiliza- and peregrine falcon. Has high injury rate and
tion delayed until after arousal from torpor. greater-than-average speed. Vocalization a high-
Gestation about 60 days with litter size 1 in W pitched, metallic squeak audible to the human
North America and 2 in Cuba and E U.S. ear (Watkins 1977).
(Kurta and Baker 1990). STATUS.—Species of special concern in
ECOLOGY.—More abundant in deciduous Utah due to declining populations (Kimball
forests and more forested, dry regions in W 1997).
U.S. In mountainous regions, males occur at
higher elevations than females. Observed leav- Idionycteris phyllotis (Allen 1916)
ing roosting site when temperature exceeds Allen’s Big-eared Bat
33–35°C. Forages at night with no preference Corynorhinus phyllotis G. M. Allen, 1916. Bull. Mus. Comp.
for over-water feeding sites. Generalist feeder Zool., 60:352. Type locality Mexico, San Luis Potosi
and has opportunistic predators. Post-sunset (Wilson and Reeder 1993).
glow and spatial memory might be relied on
for finding foraging grounds. Isolation and GENERAL CHARACTERISTICS.—Large ears
with lobes able to fold into ram’s horn fashion
feeding ultrasonic calls utilized. May also
for protection (Sparks 1974). Pelage soft black-
identify prey by low-frequency flight sounds
ish with yellowish tips (Czaplewski 1983).
of prey (Kurta and Baker 1990).
Total length: 103–121; length of tail: 46–55;
SPECIMENS EXAMINED.—Garfield Co.: 5 mi
length of hind foot: 9–12; length of ear: 38–43;
W Escalante, 3 (Hasenyager 1980); Kane Co.:
length of forearm: 43–49; body mass: 10–12 g
Kaiparowits Plateau, 2 (Hasenyager 1980).
( Jameson and Peeters 1988, Fitzgerald et al.
ADDITIONAL RECORDS.—Kane Co.: Fin Lit-
1994).
tle Spring, 5 (BLM); Pete’s Cove, 3 (BLM);
DISTRIBUTION.—Mountainous regions of
Pink Cove, 12 (BLM); Nipple Spring, 1 (BLM); SW U.S. and Mexico (Czaplewski 1983). Dis-
Drip Tank Spring, 11 (BLM); Paria River #2 tributed in Utah in southernmost counties
Site, 10 (BLM). (Hasenyager 1980, Mollhagen and Bogan 1997).
Euderma maculatum (Allen, 1892) REPRODUCTION.—Single young born in
maternity colonies between June and August
Spotted Bat (Czaplewski 1983).
Histortius maculatus J. A. Allen, 1892. Bull. Am. Mus. Nat. ECOLOGY.—Primarily dwells in forested
Hist., 3:195. Type locality Santa Clara Valley, Casrac mountains from pine, fir, and oak to more
Creek mouth, Los Angeles Co., California (Wilson and
Reeder 1993). riparian woodlands of sycamore, cottonwood,
willow, and walnut. In Utah, also found in
GENERAL CHARACTERISTICS.—White spots drier habitats. Roosts in rocky cliffs, outcrop-
on black body. Large ears. Total length: 107– pings, boulder piles, or lava flows. Food con-
115; length of tail: 47–50; length of hind foot: sists of moths, soldier beetles, dung beetles,
not given; length of ear: 44–50; length of fore- leaf beetles, roaches, and flying ants gleaned
arm: 48–51; body mass: 9–11 g (Watkins 1977). from vegetation or pursued in flight. Variable
DISTRIBUTION.—Rocky Mountain range from flying capabilities and speeds for adaptive for-
S Montana to S Mexico ranging from desert aging. Emits “peeps” similar to Euderma mac-
rubble to coniferous montane forests (Watkins ulatum but lower in pitch. Echolocation has
1977). Found in S portions of Utah (Hasenyager large range of signal types used by nearly all
1980). other bat species, allowing for versatility in
REPRODUCTION.—Fertilization occurs in orientation sounds (Czaplewski 1983).
summer months. Single offspring born mid- to ADDITIONAL RECORDS.—Kane Co.: Pink
late summer (Watkins 1977). Cove, 1 (BLM); Pete’s Cove, 2 (BLM).14 MONOGRAPHS OF THE WESTERN NORTH AMERICAN NATURALIST [No. 1
STATUS.—Registered as a species of special GENERAL CHARACTERISTICS.—Male color-
concern in Utah due to limited distribution ing bright red to orange-red; females dull red
(Kimball 1997). or chestnut; both with frost-colored back and
breast. Characterized by white patch on each
Lasionycteris noctivagans
shoulder. Ears small and rounded with small
(Le Conte, 1831)
tragus (Shump and Shump 1982a). Total length:
Silver-haired Bat 107–128; length of tail: 40–60; length of hind
Vespertilio noctivagans Le Conte, 1831. In: McMurtrie, foot: 8–11; length of ear: 8–11; length of fore-
Anim. Kingdom, 1(App.):431. Type locality eastern arm: 35–46; body mass: 7–16 g (Fitzgerald et
United States (Wilson and Reeder 1993). al. 1994).
DISTRIBUTION.—S Canada through U.S.,
GENERAL CHARACTERISTICS.—Medium-sized
Mexico, Central America, and South America
bat. Pelage nearly black with silver-tipped dorsal
to Chile and Argentina (Shump and Shump
hair. Interfemoral membrane slightly furred.
1982a). Utah records indicate presence in S
Short, rounded, naked ears (Kunz 1982). Total
two-thirds of state (Durrant 1952).
length: 90–112; length of tail: 35–48; length of
REPRODUCTION.—Breeding August–Septem-
hind foot: 8–11; length of ear: 13–16; length of
ber with fertilization in spring. Gestation 80–
forearm: 37–44; body mass: 7–15 g (Fitzgerald
90 days with 3–5 young born in June–July
et al. 1994).
(Shump and Shump 1982a).
DISTRIBUTION.—S Canada, SE Alaska, and
ECOLOGY.—Wide range of habitats used,
U.S. except southernmost regions, NE Mexico,
preferring riparian areas where roosts avail-
Bermuda (Wilson and Reeder 1993). Likely
able in tree foliage (Schmidly 1991). Solitary
statewide distribution for Utah (Hasenyager
bat. Foraging begins 1–2 hours after sunset,
1980).
feeding mostly on moths, crickets, flies, beetles,
REPRODUCTION.—Mating in temperate zones
cicadas, and grain moths. Highly migratory.
likely in autumn, with sperm storage over
Predators include falcons, hawks, opossums,
winter. Fifty- to 60-day gestation in May to
domestic cats, owls, and roadrunners (Shump
June. Two young born in early summer. Un-
and Shump 1982a).
known if species forms maternity colonies
STATUS.—Considered sensitive species in
(Kunz 1982).
Utah due to limited distribution and declining
ECOLOGY.—Found in riparian and coniferous
populations (Kimball 1997).
woodlands near streams and ponds (Hasen-
yager 1980). Thought to be solitary. Tree- Lasiurus cinereus
rooster, but utilizes mines, caves, tree hollows (Palisot de Beauvois, 1796)
and crevices in winter. Food habits opportunis-
Hoary Bat
tically insectivorous. Forages comparatively in
coniferous and deciduous forests adjacent to Vespertilio cinereus Palisot de Beauvois, 1796. Cat. Ras-
ionne Mus. Peale Philadelphia, p. 18. Type from Phila-
streams or ponds in bimodal activity of 2–4 delphia, Pennsylvania (Wilson and Reeder 1993).
hours before sunset then 4–6 hours after sun-
set. Probable that presence of red and hoary GENERAL CHARACTERISTICS.—Larger bat.
bats alters foraging time (Kunz 1982). Pale brown above with heavily frost-tipped
SPECIMENS EXAMINED.—Kane Co.: 42 mi S, pelage (Shump and Shump 1982b). Small,
18 mi W, Paria River, 1 (USNM). rounded ears and interfemoral membrane well
ADDITIONAL RECORDS.—Kane Co.: Water furred. Total length: 120–145; length of tail:
Tank Springs, Kaiparowits Plateau, 50 mi SE 49–60; length of hind foot: 9–12; length of ear:
Escalante (Tanner 1940); Tommy Water, 3 9–14; length of forearm: 46–56; body mass:
(BLM); Drip Tank Spring, 7 (BLM); Paria 18–32 g ( Jameson and Peeters 1988, Fitzger-
River #2 Site, 1 (BLM). ald et al. 1994).
DISTRIBUTION.—Covers the U.S. and Canada
Lasiurus borealis (Muller, 1776) to Guatemala, and South America from Brazil
Western Red Bat to Argentina and Chile in desert canyons,
Vespertilio borealis Muller, 1776. Linne’s Vollstand. Natur.
coniferous and deciduous forests (Shump and
System, Suppl., p. 20. Type locality New York (Wilson Shump 1982b). Statewide distribution in Utah
and Reeder 1993). (Durrant 1952).2002] MAMMALS OF GSENM 15
REPRODUCTION.—Breeding in fall and per- Myotis ciliolabrum
haps winter. Two young born in late May to (Merriam, 1886)
early June that cling to mother in day and are Western Small-footed Myotis
left hanging on a twig or leaf while mother
feeds at night (Shump and Shump 1982b). Vespertilio ciliolabrum Merriam, 1886. Proc. Biol. Soc.
Washington, 4:2, Dec. 17. Type from a bluff on Hack-
ECOLOGY.—Found in forested areas of the berry Creek, about 1 mi from Castle Rock, near Ban-
West (Hasenyager 1980). Solitary bat that usu- ner, Trego Co., Kansas (Hall 1981).
ally roosts in tree foliage. Forage primarily
includes moths and some beetles, flies, grass- GENERAL CHARACTERISTICS.—Small bat with
hoppers, termites, dragonflies, and wasps. Has yellow-brown pelage and black face, ears,
been observed when feeding on moths to bite wings, and interfemoral membrane. Keeled
from behind, thus engulfing abdomen and calcar. Dark facial mask on some individuals.
thorax and shearing off head and wings. Occa- Total length: 75–88; length of tail: 33–42;
sionally taken by hawks or owls (Shump and length of hind foot: 5–8; length of ear: 12–15;
Shump 1982b). length of forearm: 30–35; body mass: 3.5–5.5 g
ADDITIONAL RECORDS.—Kane Co.: Drip Tank (Fitzgerald et al. 1994).
Springs, 1 (BLM). DISTRIBUTION.—Throughout all W U.S. and
across the state of Utah (Wilson and Reeder
Myotis californicus 1993, Mollhagen and Bogan 1997).
(Audubon and Bachman, 1842) REPRODUCTION.—Single young born in May
California Myotis to June in small maternity colonies. Very little
Vespertilio californicus Audubon and Bachman, 1842. J. known about their reproductive activity
Acad. Nat. Sci. Philadelphia, ser. 1, 8:285. Type locality (Schmidly 1991).
“Monterey, California, U.S.A.” (Wilson and Reeder ECOLOGY.—Widely distributed in many
1993). different western habitats. Also found near
forested areas. Small maternity colonies formed;
GENERAL CHARACTERISTICS.—Light to brown
otherwise habits generally solitary. Low flier
fur with rusty tint above and paler below.
and insectivorous, opportunistic feeder. Feed-
Dark wings, ears, and interfemoral membrane.
ing habits similar to California myotis (Schmid-
Small feet. Total length: 70–84; length of tail:
ly 1991). Various roosts, especially man-made
30–40; length of hind foot: 5.5–8.2; length of
structures. Relatively high toleration to cold
ear: 11–15; length of forearm: 29–36; body temperatures; therefore shorter hibernation
mass: 3–5 g (Fitzgerald et al. 1994). than other bats (Hasenyager 1980).
DISTRIBUTION.—W North America from REMARKS.—Myotis ciliolabrum and M. leibii
Alaskan Panhandle to Baja California and Chi- synonymized by Wilson and Reeder (1993).
apas, Mexico (Simpson 1993). Records in Utah Herein, separated based on data presented by
include counties from the S three-fourths of van Zyll de Jong (1984) and revised as M. leibii
state (Hasenyager 1980). (eastern race) and M. ciliolabrum (western
REPRODUCTION.—Delayed fertilization with race) according to diagnostic bacula size, vaulted
1 young per year; breeding in late autumn to braincase, upper incisor width, and DNA work
early spring. Maternity colonies form June to using UPGMA clustering. Best and Jennings
July (Simpson 1993). (1997), in Mammalian Species account of M.
ECOLOGY.—Inhabits desert, semidesert, leibii, also considered them separate species.
grasslands, ponderosa pine, and lower Sonoran
through transitional life zones. Rocky canyons, Myotis evotis (H. Allen, 1864)
crevices, and caves used for roosting. Forages Long-eared Myotis
before sundown (Hasenyager 1980). Feeds
mainly on lepidopteran and dipteran species, Vespertilio evotis H. Allen, 1864. Smithson. Misc. Coll.,
7:48. Type locality restricted to Monterrey, Monterrey
but will take coleopterans, trichopterans, and Co., California (Wilson and Reeder 1993).
hemipterans. Roosts alone or in small groups
(Simpson 1993). GENERAL CHARACTERISTICS.—Light brown
SPECIMENS EXAMINED.—Kane Co.: Willow to brown, glossy, long pelage. Longest ears in
Tank Spring, 1 (BYU). genera, usually black with long, tiny tragus
ADDITIONAL RECORDS.—Kane Co.: Nipple (Zeveloff 1988). Total length: 88–92; length of
Spring, 1 (BLM). tail: 41–46; length of hind foot: 8–10; length ofYou can also read
