S FaunaWest Wildlife

599.74 Consultants (Firm)
442 An ecological

F2etrs and taxononlc
1991 review of the

swift fox ( Vulpes

veloxi with

AN tl'ULUWAL

REVIEW OF THE

SWIFT FOX (Vulpes velox)

WITH SPECIAL REFERENCE TO MONTANA

January 1991

ft AM

MONTANA STATE LIBRARY

S 599.74442 F2etrs 1991 C.1
An ecological and taxonomic review of th

2 2 2004

3 0864 00072197 0

AN ECOLOGICAL AND TAXONOMIC REVIEW

OF THE SWIFT FOX (Vulpes velox)
WITH SPECIAL REFERENCE TO MONTANA

prepared by

FaunaWest Wildlife Consultants
P.O. Box 113
Boulder, MT 59632
(406) 225-3221

prepared for

Montana Department Fish, Wildlife and Parks
Montana State University Campus
Bozeman, MT 59717

8 January 1991

TABLE OF CONTENTS

SECTION PAGE

LIST OF FIGURES ii

INTRODUCTION 1

DISTRIBUTION AND STATUS 1

TAXONOMY 17

SPECIES DESCRIPTION 18

REPRODUCTION 21

DENS 21

POPULATION STRUCTURE 22

MORTALITY 23

HABITAT USE 23

HOME RANGE AND DISPERSAL 25

FOOD HABITS 25

BEHAVIOR 27

MANAGEMENT AND ECONOMIC VALUE 27

PARASITES AND DISEASES 28

PREDATION 30

SURVEY TECHNIQUES 30

REINTRODUCTION 3 3

POTENTIAL MONTANA REINTRODUCTION SITES 36

RECOMMENDATIONS 38

ACKNOWLEDGMENTS 40

LITERATURE CITED 41

APPENDIX A 50

i

LIST OF FIGURES

Figure 1. Historic range of the swift and kit foxes 2

Figure 2. Probable present range of the swift fox 4

Figure 3. Present range of the red fox 5

Figure 4. Present range of the gray wolf 6

Figure 5. Present range of the coyote 7

Figure 6. Historic and present records of swift foxes in

Montana 10

Figure 7. Recent records (X) of the swift fox in North

Dakota 11

Figure 8. Established populations (P) and recent records (X)

of the swift fox in South Dakota 13

Figure 9. Current range of the swift fox in Wyoming 14

Figure 10. General release sites of swift foxes in Alberta

and Saskatchewan 15

Figure 11. Lateral, ventral, and dorsal views of a swift fox
skull showing condylobasal measurement of 105-120

mm. (Modified from Egoscue (1979)) 19

Figure 12. Physical characteristics of the swift fox 2 0

Figure 13. Tracks, trail, and scat of the swift fox.

(Modified from Murie (1975)) 32

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INTRODUCTION

The swift fox (Vulpes velox) and its close relative, the kit
fox (V. macrotis) , represent the smallest North American members of
the family Canidae. Weighing from 4 to 6 pounds and measuring just
over 2 feet, the swift fox is a uniform buffy tan except for a
black-tipped tail and dark patches on either side of its nose.
The swift fox is a predator of small mammals, birds, and insects.

The swift fox is believed to have originally been abundant
throughout its range on the Great Plains. It was extirpated early
in this century from the northern portion of its range while
remnant populations in the southern portion survived human
settlement of the prairies. The original cause of this widespread
extirpation is attributed to the ready acceptance of poisoned baits
and traps placed for coyote (Canis latrans) and wolf (Canis lupus) ,
extermination. Loss of habitat to dry land agriculture, a changing
prey base, and increased interspecific competition from coyotes and
red foxes (V. vulpes) may have also contributed to this decline.

The loss of the swift fox over such a broad area prior to the
advent of quantitative ecological studies has resulted in a paucity
of ecological information on this species. Virtually nothing is
known about this species in Montana. Its small size and nocturnal
habits have also contributed to this lack of even basic biological
information. In recent years, swift foxes have increased in
numbers and records of dispersing swift foxes have been reported in
formerly unoccupied areas. As a result, there has been increased
interest by state and provincial conservation agencies with regard
to swift fox ecology and conservation.

The purpose of this paper is to review published information
on the distribution, taxonomy, biology, ecological relationships,
and food habits of the swift fox. Conservation issues and swift
fox reintroduction programs are also reviewed. Special effort has
been placed on reconstructing the original distribution of the
swift fox in Montana, its prey base, and habitat use. This paper
is designed to provide baseline biological information on the swift
fox to aid in designing a cost effective and biologically sound
swift fox reintroduction/management program for the state of
Montana .

DISTRIBUTION AND STATUS

The swift fox originally occurred throughout the Great Plains
from the southern portion of the Canadian prairie provinces south
to central Texas (Figure 1) (Hall 1981) . The swift fox ranged as
far east as western Iowa and Minnesota, although there are no
specimen records for verification (Allen 1870, Swanson et al. 1945,
Bowles 1975) . The Rocky Mountains represent the western margin of

1

Figure 1. Historic range of the swift and kit foxes.

2

its range. Hall and Kelson (1959) show the swift fox distribution
crossing the mountains in northwest Montana and extending into
southeast British Columbia but this is without factual basis
(Hoffmann et al. 1969) and Soper (1964) does not list the swift fox
as part of British Columbian mammalian fauna. The southwestern
margin of the swift fox range in New Mexico and Texas abuts with
the kit fox range (Hall and Kelson 1959, Rohwer and Kilgore 1973) .
The present range of the swift fox is much reduced from its
historic range (Figure 2) .

The red fox now occurs in many areas formerly occupied by the
swift fox (Samuel and Nelson 1982) (Figure 3) . There is some
guestion whether the red fox is native to North America (Samuel and
Nelson 1982) , although Churcher (1959) believes it was native north
of latitude 40° N. Red foxes are apparently native to Montana
based on observations of them along the Missouri River by Lewis and
Clark in 1805 (Burroughs 1961) . Hoffmann et al. (1969) document
the rapid increase in records of this species in Montana during the
mid-1900 's. The original red fox distribution is confounded by
numerous introductions for sporting purposes as early as 1750
(Godin 1977) . Existing swift fox populations tend to occur in
geographic areas not inhabited by red foxes. A similar pattern is
also present for kit foxes.

Other members of Canidae found within the historic swift fox
range include the gray wolf, coyote, and gray fox (Urocyon
cinereoargenteus) . The former two species originally overlapped
the swift fox in both range distribution and habitat use (Figures
4 and 5) . Both of these species had larger ranges and occupied a
greater diversity of habitats. The wolf has been eliminated from
all areas of the present swift fox range. The coyote, on the other
hand, has increased in numbers to partially fill the niche
vacated by the wolf. The coyote does not compete well with the
wolf (Mech 1966, 1974) and does not tolerate foxes (Vulpes sp. and
Urocyon cinereoargenteus ) (Young 1951, Deckker 1983, Voigt and
Earle 1983) .

Despite its nocturnal habits, the swift fox was considered
common by naturalists and explorers in most areas of its original
range (Coues 1893) . Lewis and Clark reported observations of the
swift fox along the Missouri River in Montana in 1805 (Burroughs
1961) . Specific sightings were at the Great Falls of the Missouri
and Two Medicine River above the mouth of the Marias River. Lewis'
notes indicate that he had not seen the swift fox before the Great
Falls. Bailey and Bailey (1918) reported the swift fox to be
common over the plains along the eastern edge of Glacier National
Park. Although no positive record existed, they thought the swift
fox entered the Park at St. Mary's Lake and in the lower portions
of the Swiftcurrent and Belly River Valleys.

One indicator of the original abundance of the swift fox comes
from fur returns compiled by the American Fur Company and the

3

Figure 2. Probable present range of the swift fox.

4

Figure 3. Present range of the red fox.

5

Figure 4. Present range of the gray wolf.

6

Figure 5. Present range of the coyote.

7

Hudson Bay Company. Since the early 1800' s, large numbers of furs
have been traded annually. In the journals of Alexander Henry
(Reid and Gannon 1928) , 237 swift foxes were reported taken between
1800 - 1806 in northeastern North Dakota. Between 1835 and 1838,
the American Fur Company traded 10,614 swift fox pelts at their
upper Missouri and Sioux outfits (areas covering Montana and the
Dakotas) while only 1989 red fox and 108 gray fox furs were traded
(Johnson 1969) . Grinnell (1914) comments that in addition to
poisoning 750 wolves and 250 coyotes in Kansas during the winter of
1860 - 1861, several bales of yellow fox (V. velox) furs worth
$0.25 each were also taken. In the mid 1800* s an average of 5,000
swift fox pelts per year over a 24 year period were traded in
Canada (Rand 1948). As late as 1905, 5,000 "kits" were sold in a
London auction (Seton 1929) . In Canada, commercial trapping of
swift foxes continued into the early 19 00's. However, by 192 5,
only 508 pelts were sold from the Canadian prairie provinces
(Carbyn and Killaby 1989) and after this date no records were kept
on the species due to their insignificant role in the annual take
of furs in Canada.

The swift fox was extirpated from vast areas of its range
early in this century and was not considered abundant even in areas
where it survived. Causes of the decline are attributed primarily
to poison baits set for wolf and coyote control/extermination, and
the ease with which swift foxes can be trapped. Choate (pers.
comm.) now thinks that the early canid extermination efforts which
eliminated the wolf and favored the coyote resulted in biotic
communities with too much niche overlap between the swift fox and
coyote. Robinson (1953) and (1961) and Linhart and Robinson (1972)
present some data to suggest an inverse relationship between
coyotes and kit/swift foxes. Coyotes are also a known predator of
swift foxes (Herrero et al. 1990, Kilgore pers. comm., Choate pers.
comm.) and are capable of causing substantial yearly mortality
among swift fox populations. Also contributing to the decline in
swift foxes was the influx of humans to the Great Plains and the
conversion of native prairies to croplands.

The last of the original swift foxes recorded in Montana were
those reported by Bailey and Bailey (1918) . We sent letters of
inquiries to 30 of the largest museums in the United States and
Canada specifically requesting information on swift fox specimens
collected in Montana. Our intent was to establish a historical
distribution for the swift fox in Montana. Twenty-three responses
(the American Museum of Natural History with 254,800 specimens -
second largest in North America - did not respond to our inquiry)
representing a survey of 1,764,590 mammal specimens (this figure
includes collections at Montana's two universities) resulted in
locating 44 swift fox specimens collected prior to 1910. (Yates et
al. (1987) lists the total North American mammal collection at
2,823,129 specimens.) Montana swift fox specimens represent only
0.0025% of all mammal specimens surveyed. All located specimens
belong to the National Museum of Natural History (two are on

s

indefinite loan - one of these resides at the University of
Montana) . Forty-three specimens were collected at Kipp and
Blackfoot in upper Teton County, (now Glacier County) Montana
between 1901 and 1906 for the National Museum of Natural History.
One other specimen of a Montana swift fox, taken near Fort Benton
in the late 1880's (USNH A04213) , is housed in this same museum.
Appendix A lists all known Montana swift fox specimens.

The lack of any confirmed records of swift foxes in Montana
following Bailey and Bailey's (1918) report, prompted Hoffmann et
al. (1969) to declare this species extinct in Montana. They based
this conclusion on the absence of swift foxes from 16 years of fur
harvest data. This trend continued until 1978 when a male swift
fox was trapped in Custer County near the Tongue River on 5 March
1978 (T2N, R45E, Sec. 12) (Moore and Martin 1980) . This specimen
is now in the Montana State University vertebrate museum (#624 0) .
In mid-January 1984, a female swift fox was captured north of
Broadus (Henckel 1984) . Little is known about this specimen other
than it was shown to a taxidermist and was not surrendered to
Montana Dept. of Fish, Wildlife and Parks as required by law. Its
disposition and information concerning the exact location of
capture are not known. Later in the same year, another specimen
was captured on 15 November 1984 (T15N, R58E, Sec. 2) east of
Glendive in Dawson County (Vallard 1985) . This specimen was turned
over to Montana Dept. Fish, Wildlife and Parks and later sent to a
taxidermist to be mounted as a display specimen. The disposition
of this specimen is unknown at present. Pacific Hide and Fur
reported one swift fox fur traded at Havre in 1985-86 (Carbyn and
Killaby 1989) , this being caught along Sage Creek near Gilford,
Montana. Additionally, during the same period, there was one
sighting and one unconfirmed capture of a swift fox near Hogeland
and Chinook, respectively. Presumably all three of these
observations are of animals dispersing from the Canadian
reintroduction program (Carbyn and Killaby 1989) . At present, one
radio-marked Canadian swift fox is living in this area of Montana
(Dood pers. comm.). Figure 6 shows historic and recent swift fox
records and observations for Montana. The three unconfirmed
observations in southeast Montana are from Thompson (1982) .

A similar trend has been noted in other states. No sightings
of swift foxes were reported in North Dakota from 1915 to 1970
(Pfieffer and Hibbard 1970). Bailey's (1926) last record of a
swift fox came in 1915 from northern Mckenzie County, North Dakota,
where they were considered very scarce. The adult male specimen
reported by Pfieffer and Hibbard (1970) was from southeastern Slope
County, North Dakota about 161 km southeast of Glendive, Montana
(Figure 7) . Seabloom et al. (1978) report a swift fox sighting in
southern Mercer County, North Dakota, in 1976. More recently, a
swift fox was observed in May 1990 in northern Golden Valley
County, North Dakota about 80 km east of Glendive, Montana (USFWS
1990) .

9

10

11

The last record of swift foxes in South Dakota was reported by
Visher (1914) in Harding County where they were considered rare.
There was a hiatus of 52 years until the next report of a swift fox
in 1966 (McDaniel 1976) . Additional records were reported by Van
Ballenberghe (1975) which included two individuals from Fall River
County in 1970, one animal from Tripp County in 1972, and another
individual from Sulley County in 1974 (Figure 8) . Subsequently,
Sharps (1977) reported three established swift fox populations in
South Dakota (Shannon, Haakon, and Perkins Counties) . Hillman and
Sharps (1978) provide additional data on the Shannon County
population.

Likewise, swift foxes were not reported in Wyoming for many
years (Long and Long 1964, Long 1965) until the Longs' report of
one from Laramie county in 1958. Muchmore (1975), Floyd and
Stromberg (1981) , and Floyd (1983) provide additional details of
swift fox records in this area and assume continuity between the
1958 record and their earliest record of 1976. Figure 9 shows the
current range of the swift fox in Wyoming.

In Nebraska, no records of swift foxes exist from 1901 to 1953
(Jones 19 64) when a female with two kits was taken. Bluss et al.
(1967) report on a female swift fox taken in March 1966 in
McPherson County, Nebraska. Hines (198 0) presents a range map for
Nebraska showing all verified swift fox sightings since 1950 and
indicates that swift foxes range throughout most of western
Nebraska.

The swift fox also vanished from its range in Canada. The
last confirmed records of swift foxes in Saskatchewan were in 1927
and 1928 (Beck 1958) and 1938 for Alberta (Soper 1964). Swift
foxes are now present in these two provinces due to reintroduction
(Carbyn and Killaby 1989, Herrero et al. 1990) (Figure 10).
Loomand (1972) reported on a possible swift fox sighting in
Saskatchewan during the 1970' s, but no established population was
ever found.

In the southern portion of its range, the swift fox never
disappeared, although it became very rare by the 1930' s. Zumbaugh
and Choate (1985) document the historical biogeography of the swift
fox in Kansas. These authors present evidence to show the swift
fox as being extremely abundant in the mid-1800' s, declining
rapidly by the late 1800 's, becoming very rare and restricted in
distribution by the 1930 's, and a general population recovery
starting in the 1950 's (Martin and Sternberg 1955). The swift fox
is presently considered the most abundant fox of the Kansas high
plains (Zumbaugh and Choate 1985) .

In Colorado, Lechleitner (1969) reported swift fox populations
were reduced in size by 1900 and later extirpated from many areas.
Bailey (1905) considered the swift fox in Texas to be scarcer than
in former years. Glass (1959) reported a long absence of records

12

13

Figure 9.

15

for the swift fox from the Oklahoma Panhandle until 1956 but by the
mid-1960 's the swift fox was considered abundant in this area
(Kilgore 1969) .

These records indicate that the swift fox became so scarce
throughout its range that by the mid-1900' s it was believed
extirpated in most areas. This may have been an artifact of swift
fox behavior, the nature of small isolated populations, and a lack
of interest among conservation agencies. The net result was a
series of state reports (cited above) of the swift fox returning to
former habitat after being absent for an extended period. A
careful look at these reports suggests that the swift fox was
either never totally extirpated from many areas and/or it is a good
colonizer. The timing of the reports detailing increased numbers
of swift foxes in Oklahoma, Kansas, Nebraska, Wyoming, North
Dakota, and South Dakota are so closely spaced that they suggest a
response by several swift fox populations to a common variable. We
now know that reports of swift foxes reappearing in Kansas, South
Dakota, and Wyoming could be traced to pre-existing populations in
those states (Zumbaugh and Choate 1985, Sharps 1977, and Floyd and
Stromberg 1981, respectively) .

The recovery of the swift fox starting in the 1950' s has been
attributed to a change in coyote control methods; specifically the
substitution of compound 1080 (sodium monof luoroacetate) bait
stations for strychnine and the use of "coyote getters" (Robinson
1953, Robinson 1961, Linhart and Robinson 1972). Egoscue (1956)
concluded that these two methods of coyote control had no effect on
kit fox populations in Utah. President Nixon's ban in 1972 on the
use of chemical toxicants for predator control on Federal lands
(Executive Order 11643) was undoubtedly a positive step in swift
fox recovery. The recovery of the swift fox in many areas of its
former range may not be so much a spread from south to north, as a
general response of small, remnant populations scattered over a
broad geographic area to changes in Federal policy and social-
economic factors (Floyd and Stromberg 1981) . This would illustrate
the ability of the swift fox to exist in relative obscurity in
small isolated populations and to avoid detection unless specific
searches are made for this species.

The current distribution of the swift fox appears to be
composed of several highly disjunct populations. Scattered
populations exist from western South Dakota, south through eastern
Wyoming and western Nebraska, into eastern Colorado and western
Kansas, and on into New Mexico and Texas (Figure 2) . Three known
populations lie very close to Montana's borders. One population is
located southwest of Gillette/southeast of Buffalo, Wyoming
(Pallister, Boyce, and Stromberg pers. comm.) (Figure 9). The 1978
Montana record (Moore and Martin 1982) may have been a dispersing
individual from this population. Reintroduced swift foxes are now
present in southeastern Alberta and southwestern Saskatchewan
(Carbyn and Killaby 1989, Herrero et al. 1990) (Figure 10) and

16

represent the closest known population to Montana. A few of these
animals have turned up in Montana (Carbyn and Killaby 1989, Dood
pers. comm.) . An existing population of swift foxes in Perkins
County, South Dakota, is another population very close to Montana
(Figure 8) and may account for the two 1984 Montana records.

TAXONOMY

The first record of a swift fox comes from Alexander Henry's
report of fur shipments from Pembina Post of the Northwest
Company's Red River District in 1801. By 1805, 117 swift fox furs
had been shipped (Burroughs 1961) . Lewis indicates in his notes on
July 18 05 that the term "kit fox" was already in use by the fur
traders of that period. However, it was not until 1823 that the
swift fox was first described as a species by Thomas Say (Say 1823,
Burroughs 1961) . It is generally believed that the swift and kit
fox are two separate species. So similar are the two species,
however, that their common names have been used interchangeably
(Miller and Kellogg 1955) , and there has been considerable
controversy over the specific status of the two species. Rohwer
and Kilgore (1973) analyzed swift and kit fox specimens collected
along, and on either side of, a zone of sympatry in eastern New
Mexico and south-central Texas. These authors documented
hybridization between the two species in the zone of overlap but
felt that interspecific competition prevented either species from
expanding into the other's range. Others (Creel and Thornton 1971,
Snow 1973, Thornton and Creel 1975, McGrew 1979, Stromberg and
Boyce 198 6) present evidence supporting the swift fox and kit fox
species designation.

Principle differences between the two species are longer ears
and tail in the kit fox, and a larger body size and broader skull
(Egoscue 1979) for the swift fox.

Merriam, an early taxonomist prone to excessive subspecies
designations, described two subspecies of the swift fox, the
northern swift fox (Vulpes velox hebes) , and the southern swift fox
(Vulpes velox velox) (Merriam 1902) . This classification was the
basis of a brief listing of the northern subspecies as an
endangered species (U.S. Fish and Wildlife Service 1979 and 1982).
The northern swift fox was delisted when it was decided that valid
subspecies variation did not exist. Stromberg and Boyce (198 6)
examined swift fox specimens collected from throughout their range
and concluded that subspecific status for a northern population as
proposed by Merriam (1902) is not justified. However, these
authors cautioned that there was significant geographic variation
among the specimens examined and that this variation may reflect
genetic differences. They advised that conservation efforts to
restore swift foxes to former portions of their range consider this
geographic variation as a preservation criteria. A similar study
conducted by Dragoo et al. (1987) also concluded that the

17

subspecific designation as proposed by Merriam (1902) was not
valid.

Hall (1981) suggests that the swift and kit foxes are
conspecifics and may be only subspecif ically distinct. Dragoo et
al. (1990) present data to support this contention. They assessed
the relationships of these two foxes by morphometric and protein-
electrophoretic methods. In the latter case, they found genetic
divergence to be negligible with a high degree of genetic
similarity among all nominal subspecies examined. Morphometric
analysis were only able to distinguish between the swift and kit
fox and not between any of the previously proposed subspecies.
Dragoo et al. (1990) propose to reclassify the swift and kit foxes
as a single species - Vulpes velox - with only two recognized
subspecies - the swift fox, Vulpes velox velox. and the kit fox,
Vulpes velox macrotis.

Although the red fox (third North American member of the genus
Vulpes) appears quite distinct from the swift and kit foxes,
Thornton et al. (1971) and Creel and Thornton (1974) have
documented hybridization between kit and red foxes in Texas. Based
on this finding, and the report by Rohwer and Kilgore (197 3) , the
possibility appears to exist for hybridization between the swift
and red fox as well.

SPECIES DESCRIPTION

The swift fox averages about 2.4 kg body weight (range = 1.8
to 3.0 kg) and measures 70 - 85 cm for total length (Kilgore 1969).
Tail length is 24 - 35 cm, hindfoot 11 - 14 cm, and ear length 56 -
75 mm. Although males tend to be 8% heavier than females (Egoscue
1979) , skeletal sizes of the two sexes are similar (Dragoo et al.
1987) . The cranium and rostrum of a swift fox skull is relatively
short and broad (Figure 11) (condylobasal length 105-120 mm) which
distinguishes it from the longer narrower red fox skull
(condylobasal length 120-150 mm) . The lyre-shaped sagittal crest
of the gray fox distinguishes this species from the members of the
genus Vulpes. The dental formula for the swift fox is i 3/3, c
1/1, p 4/4, m 2/3.

Pelage is a buffy tan above with a light tan to white below.
A tinge of orange may be present on the sides, legs, and lower
surface of the tail. The tail has a black tip and a dark patch is
found on either side of the muzzle. Figure 12 shows the physical
characteristics of the swift fox. The swift and kit foxes are
distinguished from other North American foxes by their small size,
black tip to the tail and tan fur (Figure 12) . The foot pads of
the kit and swift foxes are largely hidden by fur; some (Seton
1929) have speculated that this aids in travel over sandy country.
Summer fur is short and coarse and winter fur is long, dense, and

18

Figure 11. Lateral, ventral, and dorsal views of a swift fox skull
showing condylobasal measurement of 105 - 12 0 mm. (Modified from
Egoscue (1979) ) .

19

small size (total length <80 cm)

lighter in color. A full account of pelage characteristics is
described by Audubon and Bachman (1854), Baird (1857), and Bailey
(1926, 1931).

REPRODUCTION

Timing of breeding in the swift fox varies across its range.
The swift fox breeds in December and early January in Oklahoma
(Kilgore 1969), January and February in Colorado (Fitzgerald et al.

1983) , and in February and early March in Nebraska (Hines 1980) .
The swift fox is believed to be monogamous (Kilgore 1969) , however,
Kilgore (1969) and Hines (1980) site examples of two males present
at a den site during the breeding season. The swift fox is
monoestrous (Asdell 1964) . The gestation period is 52 - 53 days
(Sharps 1984, Sharps and Whitcher 1984), which is slightly longer
than the 49 - 50 days for red foxes (Asdell 1964) . Young are born
in March and early April in the south, and April and early May in
the north. Eyes and ears of young open at about 10 - 15 days.
Young emerge from dens at three to four weeks of age (Hines 1980)
and are weaned at 6 - 7 weeks (Kilgore 1969, Hines 1980). Pups
weigh less than a 1/4 kg at birth but grow rapidly and obtain adult
weight by early fall (Kilgore 1969) . Dispersal of pups does not
occur until late summer or early fall. Swift and kit fox pups stay
with family groups and use their natal dens longer than most other
North American canids.

Swift foxes are capable of breeding during their first year
(Sharps 1984) , but under natural conditions in Kansas only 10% of
a sample of 100 juveniles showed evidence of breeding (Zumbaugh

1984) . In the same study, 76% of the yearling females bred and 86%
of the older females bred. Mean number of placental scars per
pregnant female in this study was 2.3 for juveniles and 4.3 for
yearlings and older. Other reports of litter size appear to be
based on number of young observed at the den. Kilgore (19 69)
summarizes his data with other accounts (Seton 1929, Cutter 1958a,
Glass 1956) of litter size (N = 9) and reports an average litter
size of 4.8. Hines (1980) reports an average litter size of 4.25
for four litters and Fitzgerald et al. (1983) had an average litter
size of 3.6 at 16 natal dens. Litter size in captive-raised swift
foxes has ranged from 2-7 (Carbyn and Killaby 1989) .

DENS

Swift and kit foxes are the most den dependent members of the
North American canids (Egoscue 1979) . Dens are used on a daily
basis year long (Kilgore 1969) . Swift foxes can construct their
own dens (Kilgore 1969) , or may make use of burrows dug by prairie
dogs (Cynomys ludovicianus) or badgers (Taxidea taxus) . Uresk and
Sharps (1986) reported that swift foxes favored hilltops as den
sites. Most dens were located in short to mid-grass prairies and

21

showed no selection for soil type. Over half of the 3 5 dens found
by Kilgore (1969) were in cultivated fields or other man-made
habitats (cemeteries) . The remainder of his den sites were in
short grass pastures. Dens were located in level areas or along
blow-ridges. Dens in cultivated fields usually had only one
opening, while in native habitat they had multiple openings (up to
9, mean 4). Fitzgerald et al. (1983) found the majority of their
dens on flat areas along ridges or drainages dominated by short
grasses. Eighty-five percent of their dens had only one or two
openings. Fitzgerald et al. (1983) also reported dens to be
clustered in groups of 3 to 13 . They found natal dens to be used
in consecutive or alternate years. Cutter (1958a) also reported
dens to show a clumped distribution with three to six dens in a
quarter of a section. Approximately 3/4 of his den sites were
located in overgrazed pastures devoid of any shrubs. Other den
sites included plowed fields and fence rows with wind-formed soil
banks.

Hines (1980) gathered information on 40 dens used by radio-
marked swift foxes. Numbers of dens used by marked animals over an
extended period ranged from 2 to 11. Males and unmated females
established non-natal dens throughout the year, and mated females
established non-natal dens except during whelping. Mated females
established natal dens from May through August. Hines (1980)
reported dens within home ranges to show a clumped distribution.
About half of his dens were located in flat terrain and the
remainder in areas of gently rolling topography. All dens were
located in short grass prairies and 68% were within 2 00 m of a
road. Hillman and Sharps (1978) also comment that all of their den
sites were within 1.6 km of a road. Hines (1980) found natal dens
to average (3.1) more entrances than non-natal dens (1.4).

All accounts of excavated dens (Cutter 1958a, Kilgore 1969,
Fitzgerald et al. 1983) suggest that swift fox dens are relatively
shallow (around 1 m deep) and range in length from 2 -5 m. Burrows
used by swift foxes have been variously reported to be 13 - 20 cm
in diameter.

POPULATION STRUCTURE

Reasonable information on age and sex structure of swift fox
populations come from two studies (Zumbaugh 1984, Fitzgerald et al.
1983) . Both of these studies show that the juvenile age class
dominates. Sixty-three percent of 298 foxes taken over two
trapping seasons in Kansas were under one year of age. Yearlings
accounted for 16% of the harvest and animals 2-4 years of age
accounted for 18% of the harvest. None of the swift foxes examined
were judged to be over 5 years of age. Hines (1980) sectioned
canines and looked at dental cementum annuli for eight foxes and
found all to be under three years of age. Fitzgerald et al. (1983)

22

reported 45% of 66 swift foxes trapped and marked on his study area
as juveniles.

Sex composition of swift fox populations appear to approximate
an equal number of males and females. In Kansas, 46% of 296 swift
fox carcasses were determined to be males and 54% were females. In
Fitzgerald et al.'s (1983) study, 53% of the animals marked were
males and 47% were females.

MORTALITY

Swift fox populations are characterized by high annual
mortality. Human related activity appears to be one of the most
common causes of mortality. This includes being struck by
vehicles, shot, and trapped (Hines 1980, Sharps and Whitcher 1984
Fitzgerald et al. 1983). Coyote predation is frequently cited as
a common form of natural mortality (Kilgore 1969, Carbyn and
Killaby 1989, Herrero et al. 1990) although there is little
evidence of such. Many authors have noted the dependency of swift
foxes on dens and interpreted this as an indication of swift fox
vulnerability to predation (Egoscue 1979) * Golden eagles (Aquila
chrysaetos) and other raptors have been found to feed on swift fox
but in many cases it is not known if they had killed the animals or
if they were scavenging (Pfeifer and Hibbard 1970, Hines 1980,
Fitzgerald et al. 1983) . Sharps (1989) observed an unsuccessful
attack on a swift fox by a great-horned owl (Bubo virqinianus) .

Sharps and Whitcher (1984) radio-marked 14 swift foxes and,
over a one year period, found 50% of the marked animals dead and
were unable to account for another 27%. Hines (1980) radio- marked
seven adult swift foxes and was able to follow these up to six
months. Three of these were known to have died and the fates of
the other four were unknown at the end of the study. He also
reported 46% mortality of juvenile swift foxes within the first
four months of post emergence from the den. Fitzgerald et al.
(1983) reported a minimum of 17% mortality of marked pups within
their first year. This was based on tag returns and actual
mortality was probably much higher. Swift fox harvest data
obtained over a two year period in Kansas would indicate almost 75%
mortality among juveniles and 50% mortality among adults (Zumbaugh
1984) .

HABITAT USE

Although no quantitative analysis of swift fox habitat
selection has been undertaken, numerous studies indicate that swift
foxes use, and prefer, short to mid-grass prairies (Cutter 1958a,
Kilgore 1969, Hillman and Sharps 1978, Hines 1980, Fitzgerald et
al. 1983) . Prairie habitats dominated by buffalo grass (Buchloe
dactvloides) or blue grama grass (Bouteloua gracilis) appear to be

23

the areas where the highest densities of swift foxes are found in
the southern portion of their range (Kilgore 1969, Hillman and
Sharps 1978, Fitzgerald et al. 1983). Dense shrub layers are not
reported as being part of swift fox habitat. In fact, most studies
report few, if any, shrubs in areas used by swift foxes. However,
this may be an artifact of surviving swift fox populations being in
geographic regions devoid of shrubs. Fitzgerald et al. (1983)
found swift fox densities in their better habitat to range from 0.8
to 1.1 animals per 100 ha and to range from 0.2 to 0.4 in poorer
habitat.

The swift fox is known to inhabit areas of mixed agricultural
use. Stromberg (pers. comm.) stated that swift foxes can be found
in these areas but in lower densities. Kilgore (1969) , Fitzgerald
et al. (1983) , and Hines (1980) reported similar results. The
lower densities of swift fox in agricultural areas may be due to
increased human caused mortality in these areas. The San Joaquin
kit fox appears to be very adaptable to substantial modification of
its habitat provided the prey base remains, and adjusts to human
presence to the point of being habituated to people (personal
observations) .

The swift fox is a prairie dwelling animal and most
topographic situations reported used by this species are either
level or gently rolling hills with slopes under 15% (Hillman and
Sharps 1978, Hines 1980, Fitzgerald et al. 1983). However, Hillman
and Sharps (1978) reported swift foxes to den within 1.6 km of the
White River in South Dakota. Original accounts of swift foxes
indicate that they also frequented the footslopes of the northern
Rockies in Montana and Alberta (Bailey and Bailey 1918, Soper 1964)

Sharps and Whitcher (1984) considered the presence of water in
swift fox home ranges as an important habitat factor. Hines (1980)
found no obvious relationship of swift foxes to water sources. All
dens located by Hines were greater than 4 km from a water source.
Hines (1980) points out that studies on the red fox show their den
sites to be closely tied to water sources, usually within 1 km of
water. This factor may provide some niche separation between swift
and red foxes in areas where they coexist.

Hines (1980) located radio-marked swift foxes and dens
disproportionately close to roads. Sixty-eight percent of his dens
were located under 2 00 yards from a road. Hillman and Sharps
(1978) reported all their dens to be located less than 1.6 km from
roads. Roads and trails may be used as travel routes for swift
foxes. This may also explain the high incident of traffic
mortalities in many studies. Madson (1990) reported that Canadian
Wildlife Service personnel successfully trapped swift foxes by
setting traps along fences which served as travel ways. One factor
which may favor the use of roadsides as swift fox den sites is that
larger predators such as coyotes may avoid (or are selectively
removed from) these areas. Some other wildlife species which may

24

serve as swift fox prey are associated with roads and trails. In
Montana, prairie dogs have been noted to disperse along roads and
prairie dog colonies frequently have roads or cow trails running
through them (Knowles 1984) .

HOME RANGE AND DISPERSAL

Home range size of resident swift foxes has only been
investigated in one study in Nebraska (Hines 1980) . Additionally,
radio collars have also been placed on swift foxes in two
reintroduction programs (Herrero 1984, Sharps and Whitcher 1984,
Herrero et al. 1990) . Average home range size for seven radio-
marked swjft foxes reported by Hines (1980) was 15.22 (range 6.66 -
28.80) km . Males had larger home ranges than females (males X =
17,33, range 6.66 - 28.80 km ; females X = 12.41, range 9.14 - 14.3
km ) . Average distance traveled over 4 7 nights for the seven
radio-marked foxes was 13.06 km. Sharps and Whitcher (1984)
commen£ that swift fox home ranges during2spring ranged from 1.6 to
3.2 km and increased to 11.2 to 12.8 km during the remainder of
the year. However, details of these data are not provided in the
paper .

Juvenile swift foxes disperse during late summer and early
fall. Several studies report dispersal distances for marked
animals. Dispersal distances bear direct relevance to the recent
Montana swift fox records. Sharps and Whitcher (1984) reported
dispersal distances of reintroduced swift foxes to range from 6 to
252 km (average distance; males 51.4 km, females 76.5 km) over
approximately a two year period. This would represent a minimum
since some of the released foxes died early in the study or radio
contact was lost. Fitzgerald et al. (1983) reported dispersal
distances for two foxes at 40 and 70 km. Carbyn and Killaby (1989)
monitored the movements of a released fox that traveled 50 km
before mating and establishing a den.

FOOD HABITS

Swift fox food habits probably have been investigated more
thoroughly than other aspects of their ecology. In general, swift
foxes appear to be opportunistic feeders (Loy and Fitzgerald 1980)
taking small mammals, birds, and insects depending on availability.
Mammals are the major component of the swift fox diet and
lagomorphs are consistently represented in food habit studies
(Cutter 1958b, Kilgore 1969, Hillman and Sharps 1978, Hines 1980,
Choate and Fox 1985, and Uresk and Sharps 1986) . Lagomorphs taken
by swift foxes include the genus Lepus and Sylvilaqus . Jackrabbits
appear to be the largest prey taken by swift foxes although they
are known to scavenge on dead ungulates including domestic cattle
(Hines 1980, Uresk and Sharps 1986) . Hines (1980) considered

25

carrion to be an important component of the swift fox diet with
road-killed animals a readily available source.

Uresk and Sharps' (1986) study is the only investigation where
prairie dogs were the major prey item and rabbits did not comprise
the bulk of the diet. This can be related to the abundance of
prairie dogs on their study area and the fact that two dens were
located in prairie dog colonies. Prairie dogs appear in the swift
fox diet in other studies where prairie dogs were present on the
study area (Kilgore 1969, Hillman and Sharps 1978, Hines 1980,
Choate and Fox 1985) . Ground squirrels, primarily thirteen-lined
ground squirrels ( Spermophilus tridecemlineatus) , are also taken by
swift foxes (Cutter 1958b, Kilgore 1969, Hillman and Sharps 1978,
Hines 1980, Uresk and Sharps 1986) . The presence of prairie dogs
and ground squirrels, two strictly diurnal genera, in the swift fox
diet implies that swift foxes are capable of diurnal hunting.

Sharps (1989) was able to return to his former study area and
conduct swift fox surveys following a 92% reduction in black-tailed
prairie dog colonies. Sharps noted a wide-scale reduction of swift
fox population densities over the survey area. During 102 days of
survey time, he was unable to locate swift foxes in the remaining
prairie dog colonies. These colonies were located in rough
topographic situations which tend to favor coyotes and bobcats
(Lynx ruf us) , Preliminary observations of swift fox scats from the
survey area in 1989 indicated that small mammals, insects, and
ground roosting birds comprised the primary food base. These data
(Uresk and Sharps 1986, Sharps 1989) suggest that prairie dogs play
an important role in swift fox ecology.

Egoscue (1975) reported kit fox welfare to be tied directly to
jackrabbit populations. Declining rabbit numbers resulted in non-
breeding and smaller litter sizes among kit foxes. He stated that
kit foxes were incapable of altering their predatory habits to take
advantage of diurnal prey species. Our observations of the kit fox
in southern California and Nevada indicate that this species is
indeed highly nocturnal. In addition, Egoscue found that the lack
of rabbits in the kit fox diet was not compensated for by nocturnal
small mammals such as kangaroo rats f Dipodomvs sp.). Because of
its dependence on rabbits, Egoscue (pers. comm.) recommends that
reintroduction efforts for the swift fox be keyed to peaks of
rabbit cycles.

Swift foxes are also reported to eat a variety of mouse
species. These include deer mouse (Peromvscus maniculatus) ,
northern grasshopper mouse ( Onvcomvs leucoaaster) . Ord's kangaroo
rat (Dipodomys ordii) , pocket mice (Perocrnathus sp.) , harvest mice
( Rvthrodonomvs sp.), prairie vole (Microtus orchocrastor) . and
cotton rat, ( Siamodon hisidus) . Insects, when seasonably
available, may be an important prey item. A variety of insects may
be taken, but grasshoppers tend to dominate in the diet (Cutter
1958b, Kilgore 1969, Hines 1980). Grasses and other plants are

26

also eaten by swift foxes and compose an important portion of their
diet even in the winter (Cutter 1958b, Kilgore 1969, Uresk and
Sharps 1986, Hines 1980, Zumbaugh et al. 1985)

BEHAVIOR

Little is known of swift fox behavior. Swift foxes are
primarily nocturnal and are only rarely seen abroad during daylight
hours. This may be a contributing factor for the paucity of
behavioral information. It may also result in small, isolated
populations going undetected unless specific surveys by experienced
biologists are employed. Hines (1980) noted swift foxes to cease
their nocturnal activity period well before sunrise and not to
begin nightly activities until nearly dark. Swift fox activity
increased after sunset and reached a plateau between 0200 and 03 00
hrs. Average distance traveled during this time period ranged from
1.22 to 1.30 km per hour. Hines (1980) considered the swift fox to
rest only during the day. The activity patterns of swift foxes may
be very similar to that of the black-footed ferret (Mustela
niqripes) .

Like black-footed ferrets, young swift foxes may be seen
playing during daylight hours at den openings (Kilgore 1969,
Chambers 1978, Hines 1980). Females with pups were noted to make
short daytime forays for food (Hines 1980) . Kilgore (1969)
considered swift foxes to pair for life but lacked any hard
evidence for this statement. He cites an example of one male being
observed at a den with at least three different females (the first
two females were killed at the den) over a two month period. This
may be taken as an example of serial monogamy or a polygamous
breeding unit (Egoscue 1962) . Hines (1980) had an observation of
two males at a den with one female. Males participate in the
rearing of young swift foxes (Hines 1980) . However, information
gathered by Hines (1980) would indicate that participation by males
is variable and often minimal.

Shortly after swift foxes are weaned, the family unit may move
from the natal den to a nearby den. This may be due to a build up
of ectoparasites in the den or exhaustion of the forage base near
the den (Jon Sharps, pers comm.). Family units split up and
dispersal of pups begins in late August. At this time, pups
resemble adults in pelage, and weigh about 2/3 of the adult weight.
In addition to the lighter weight, young during the fall may be
distinguished from adults by shorter fur on the tail, and lack of
tooth wear (Hines 1980) .

MANAGEMENT AND ECONOMIC VALUE

Swift fox fur is not particularly valuable and is dyed to be
used as an imitation for more expensive fox fur (Arnold 1925) . Its

27

lack of value, however, has not stopped the trade in swift fox
furs. As a result of its low economic value and relative scarcity,
state conservation agencies have made little effort on managing
swift fox populations.

Classification of the swift fox varies by state. Wyoming and
New Mexico have no restrictions on the harvest of swift fox. The
swift fox is considered a furbearer in Colorado, Kansas, Montana,
and Texas. These states have regulated trapping seasons, although
Montana's season is closed year long. In North Dakota, South
Dakota and Nebraska, the swift fox is classified as either
threatened or endangered and are fully protected.

Kansas took advantage of a newly instigated trapping season in
the fall and winter of 1982-1983 to gather age structure, sex
ratio, and food habits information on the swift fox (Zumbaugh 1984,
Zumbaugh et al. 1985). The first two seasons of data collection
resulted in 83 and 215 swift fox carcasses being turned over for
research purposes. Hines (1980) summarizes harvest data from the
late 1970' s for four states (Wyoming, Colorado, New Mexico, and
Texas) . The greatest number of animals harvested was in Colorado
where 1499 were taken in 1978-79. Wyoming reported the smallest
harvest with their largest being 14 in 1977. The other two states
were intermediate to these extremes.

In the United States, the swift fox is presently listed as a
category II species (Fish and Wildlife Service 1989) and has no
legal Federal protection. The northern swift fox was briefly
listed as an endangered species in 1979 but was quickly delisted
when it was determined that there was no basis for the subspecies
designation. Depending on the outcome of the present debate over
taxonomic classification of the swift and kit foxes, additional
changes in Federal classification of the swift fox may be needed to
reflect the latest scientific evidence.

The swift fox is listed in Appendix 1 of the CITES agreement,
and it is listed in the IUCN red data book (Schroeder undated) .
The Committee On the Status of Endangered Wildlife In Canada lists
the swift fox in Canada as an extirpated species (Carbyn and
Killaby 1989) .

PARASITES AND DISEASES

Relatively little work has been done on the parasites and
diseases of swift fox. Swift foxes have been noted by several
researchers to host a large number of ectoparasites (Kilgore 1969,
Hillman and Sharps 1978, Hines 1980). Kilgore noted the only swift
foxes he examined to be free of fleas were three whelps less than
10 days old. Common flea species found on the swift fox include
Pulex irritans and P. simulans (Jordan and Rothschild 1908, Holland

28

1949, Rapp 1962, Kilgore 1969). Kilgore also found the flea
Qpisocrostis hirsutus . which normally infests black-tailed prairie
dogs, at swift fox dens. Kilgore reported large numbers of fleas
present in swift fox dens and suggested that flea infestations may
partially account for the frequent changes in den sites.

Kilgore (1969), Hillman and Sharps (1978), and Hines (1980)
all report ticks (Acarina) on swift foxes. Kilgore (1969) and
Hines (1980) both identified the ticks as (Ixodes kingi) . Hillman
and Sharps (1978) noted ticks to be attached to the ears of swift
foxes .

Cutter (1958b), Kilgore (1969), and Hines (1980) identified
several endoparasites. These included the Cestodes Taenia
multiceps and Dipylidium caninum and the Nematodes Toxocara canis
and Pysaloptera sp. . Stiles and Baker (1935) and Erickson (1944)
also found Ancylostoma caninum in swift foxes. In addition, three
species of parasitic protozoans have been reported in swift foxes.
These are Coccidium biqeminum (Weidman 1915) , Isopora bicrmina
(Becker 1934, Stiles and Baker 1935), and I. f elis (Stiles and
Baker 1935) . Kilgore (1969) notes that most of the ecto- and
endoparasites are also found in other wild canids and dogs.

The occurrence of infectious diseases of wild and domestic
canids in swift foxes is not well known. However, it is reasonable
to suspect that the swift fox is susceptible to most diseases which
plague other canids. Disease should not be overlooked as an
important factor limiting swift fox populations and slowing the
return of the species to unoccupied habitat.

The San Joaquin kit fox has been surveyed for selected
diseases (McCue and O'Farrell 1988). Canine parvovirus was the
major disease of the kit fox and may well be the most important
disease among wild canids (Johnson pers. comm.). McCue and
O'Farrell (1988) found antibodies against this pathogen in all
animals examined in one collection period and in 67% of the animals
tested in a later collection period. Other diseases with a high
incidence of occurrence were canine distemper virus (0 - 14%) ,
infectious canine hepatitis virus (6 - 21%) , tularemia (Francisella
tularensis) (8 - 31%) , brucellosis (Brucella abortis and B. canis)
(3 - 8% and 0 - 14%, respectively), toxoplasmosis (Toxoplasma
gondii) (6%) , and coccidioidomycosis ( Coccidiodes immitis) (3%) .

Rabies is a virus which is known to impact wild and domestic
canids and could potentially affect swift fox populations.
Fitzgerald (pers. comm.) believes rabies may influence swift fox
numbers in northeastern Colorado. Canine distemper could also
potentially impact swift fox populations in a manner similar to
that seen in black-footed ferrets (Thorne and Williams 1988) ,
although there are no documented accounts of distemper decimating
a swift fox population. Johnson (pers. comm.) considers lyme
disease as another potential disease of the swift fox.

29

PREDATION

The small size of the swift fox makes it vulnerable to other
predators. Many researchers have commented that the swift and kit
fox are the most den dependent members of the North American
canids. Escape from predation is frequently cited as the cause of
this den dependency. Wild predators of the swift fox include the
coyote, red fox, bobcat, badger, and golden eagle. Herrero et al.
(1990) states that coyotes are a major cause of death among radio-
marked reintroduced swift foxes in Alberta. Golden eagles have
been found on swift fox carcasses on at least two occasions
(Pfieffer and Hibbard 1970, Fitzgerald et al. 1983) and noted to
kill reintroduced swift foxes in Canada (Herrero et al. 1990) .
Hines (1980) reported finding unidentified raptor pellets at the
remains of a dead swift fox but the cause of death was not
determined.

Nearly all researchers comment that human related mortality is
the principal cause of death among swift foxes. Swift fox behavior
seems to contribute to their susceptibility to humans. Swift foxes
are readily trapped (Bailey 1926) , easily shot at den sites (Hines
198 0) , and occur in areas inhabited and traveled by humans (Cutter
1958a, Hines 1980) . Sharps and Whitcher (1984) found 50% annual
mortality among radio-marked captive-released and wild swift foxes.
The four documented mortalities among the captive-releases were
human caused (two road kills, one trapped, one shot) . Hines (1980)
found road kills accounted for four of the 11 mortalities he
documented, three were shot, and cause of death was not determined
for four others. Carbyn and Killaby (1988) report similar findings
for captive-released foxes in Saskatchewan. Road kills, shootings,
and trappings seem to account for most of the known mortality.
Kleiman and Brady (1978) considered canid populations with a high
juvenile proportion to be indicative of heavy human exploitation.
They theorized the juvenile segment of an unexploited canid
population should be under 50%.

SURVEY TECHNIQUES

With the increased interest in the swift fox over the past two
decades, techniques have been developed to locate and survey swift
fox populations. These methods include fur harvest surveys, scent
post and tracking surveys, spotlight surveys, and live-trapping.

Fur harvest surveys are dependent upon having an open, legal
trapping season on swift foxes. Colorado, Kansas, New Mexico,
Texas, and Wyoming have obtained distributional information on
swift fox by tallying harvests by counties. Kansas conducted a fur
harvest survey prior to, and after, an open season on swift fox
(Zumbaugh 1984) . They found that all reports of pelts traded prior
to the open season were supposedly taken in Colorado which had an

30

open season on swift fox trapping. With the initiation of an open,
legal trapping season, number of pelts reported went up and they
were all taken within Kansas. Occasionally, fur dealers may report
a swift fox record even during a closed season (Carbyn and Killaby
1989) . However, when swift foxes were afforded legal protection in
South Dakota, Sharps (pers. comm.) found fur dealers very reluctant
to disclose information concerning the take of this species.
Government trappers have proved valuable in locating small isolated
populations of kit fox in Utah (Egoscue pers. comm.).

Scent post (Linhart and Knowlton 1975) and tracking surveys
(Sharps pers. comm.) may disclose locations of swift fox
populations. Zegers (197 6) , in developing survey techniques for
swift foxes, reported that the scent post survey was the most
effective method tested, and that swift foxes were readily
attracted to a scent station. The scent post survey may provide
presence/absence data on the swift fox and give some indication of
abundance relative to other wild canids. Muchmore (1975) ,
reporting on the results of two scent post surveys conducted in an
area of southeast Wyoming where a swift fox was observed in 1972,
stated that there were 13 swift fox visits, 23 red fox visits, and
41 coyote visits at 100 stations over a 5 day period. Results of
this survey may indicate that these three canid species are capable
of coexisting in a limited geographic area. Scent post surveys
have also been successfully used to census kit foxes in Oregon (D.
Edge pers. comm.)

Sharps (pers. comm.) stated that looking for swift fox tracks
around stock ponds, on cattle trails, or during periods of winter
snow cover, may also be useful. Pallister (pers. comm.) cautioned
that swift fox tracks may easily be mistaken for jackrabbit tracks.
Snow-tracking techniques developed for black-footed ferret surveys
(Richardson et al. 1984) may be a useful method of locating swift
fox populations. Both these techniques require that field
personnel have prior experience in discerning track sign of red
fox, gray fox, and coyotes from swift fox. Separation of sign of
these species can be accomplished with some training and practice
(Sharps pers. comm.). The tracks, trail and scat of the swift fox
is illustrated in Figure 13. This technique represents a low cost
survey method which can be applied to a broad region.

Spotlight surveys are also useful in locating and observing
the primarily nocturnal swift fox. Spotlighting as a technique
developed for locating black-footed ferrets (Campbell et al. 1985)
may be applied to locating swift fox populations. Hines (1980)
reported swift fox dens to be located disproportionately close to
roads which facilitates this technique. The habitat preference of
the swift fox for short and mid-grass prairies also makes this
technique highly useful. Hillman and Sharps (1978) used this
technique to locate swift foxes on their study area. Spotlighting
is conducted from a vehicle while driving roads and trails at night
and using a 200,000 candle power spotlight for illumination. Swift

31

64 mm

38mm

< >

Hind

32 mm

Figure 13. Tracks, trail and scat of the swift fox. (Modified
from Murie (1975)).

32

fox eyes reflect a blue-white light (Sharps and Whitcher 1984) . An
advantage of this technique is that an observation of a swift fox
results in a positive record and little prior experience is needed
to identify a swift fox. Lack of a swift fox sighting using this
technique does not mean swift foxes are absent, because only a
small area either side of a road is sampled. Sharps (pers. comm.)
considered late May to late July as the best time to conduct
spotlight surveys for the swift fox. Swift fox family units were
present around their dens at this time and were highly visible.
Foxes may be pressured into revealing their den site at night by
following them with a vehicle. If this cannot be done, the
location of the observation can be flagged and a 0.5 to 1 km area
around the observation site can be intensively searched the next
day. Spotlighting during the fall dispersal may also be a useful
time period to make observations.

Live-trapping can also be used as a sample method (Stromberg
pers. comm.). Stromberg recommended spacing live-traps at 1.6 km
intervals and baiting with bacon. Live-traps combine the greater
survey area of scent stations, and the positive record value of
spotlight surveys. However, live traps are expensive, hard to
transport, and require monitoring. Absence of a swift fox capture
can be considered as lack of swift foxes in the area (Stromberg
pers . comm. ) .

REINTRODUCTION

Reintroduction of swift foxes as a means to hasten recovery of
extirpated populations has been attempted in two separate areas.
The largest and longest running reintroduction program is in
Canada. A total of 284 swift foxes have been released in the
southern prairies of Alberta and Saskatchewan with releases
occurring every year since 1983 (Herrero 1984, Carbyn and Killaby
1989, Herrero et al. 1990). South Dakota conducted the first, but
smaller, reintroduction program with the releases taking place in
1980. The initial Canadian effort was patterned after the South
Dakota release.

Two basic methods of release have been employed in these
reintroduction efforts. The first method used was termed a "soft
release" where swift foxes are paired and held in a pen at the
release site for an extended period prior to release. In this
method, foxes must be supplied with food and water while held
captive. The pair may be held on the site long enough to breed and
rear pups. After the release, pens are kept in place and are
available to the foxes as a den site. In some cases, food is made
available at the pen following the release. Many of the animals
used in this method have been born and raised in captivity. This
method is very labor intensive, requires considerable materials,
and has a high cost per fox released.

33

The other method used for reintroduction is the "hard release"
where swift foxes are captured from existing populations in one
area, transported to the reintroduction site and released. The
animals are handled as little as possible and held a minimum of
time with this technique. Labor and material costs per fox
released are considerably lower with this technique, but foxes may
not stay at the release site.

Survival rate of reintroduced foxes has been low. Sharps and
Whitcher (1984) used both captive-raised and wild-trapped foxes for
their reintroduction program. Known mortality was 67% for the
three captive raised foxes and 50% for the five wild-trapped foxes
(fate of one was not determined) within the first year of release.
Only one pair remained in the release area indefinitely and raised
eight pups the following spring. Both individuals of this pair
were wild-trapped. All captive-born and captive-raised animals
left the release area within one month, and one moved 203 km before
establishing a home range. Sharps and Whitcher (1984) used
juvenile foxes for their release and held them in pens up to 1.5
months .

The Canadian reintroduction effort has employed both soft and
hard releases. Animals for the soft releases have been both
captive-raised and wild-trapped, while those for the hard releases
were wild-trapped. Animals used for the soft releases have
generally been paired and placed in pens at the release site during
winter and released the following spring or summer. In some cases,
the pair bred and pups were raised and also released. In the
Alberta program, animals were fed at the pen site even after the
pens were open. In the Saskatchewan program, animals were not
supplied with supplemental food after they were released.

Three of the 17 soft release foxes released in 1984 in
Saskatchewan died within 3 weeks of release. It is believed that
starvation contributed to their death (Carbyn and Killaby 1989) .
Road kills, trapping, and shooting were other documented
mortalities. By the spring of 1987, only 22% of the 39 soft
released swift foxes were known to still be alive. Initial
survival of the hard release animals appears to be better than the
soft release, however, record mild winter weather may have
contributed to this higher survival rate (Carbyn and Killaby 1989) .
Some reintroduced foxes were known to over-winter and produce pups.

Of the 25 foxes initially released in Alberta in September
1984, 17 were radio-marked, and seven of these died within 9
months, three were missing and seven were grouped in three pairs.
Each pair produced a litter (5, 4, and 3) even though they were
only one year old. Predation by coyotes and bobcats were the
causes of death in six of the seven confirmed mortalities (Herrero
pers . comm. ) .

34

In the Alberta reintroduction project, the majority (52 of 83)
of the soft released foxes stayed in the vicinity of their pens
(Herrero et al. 1990) . In fact, 52% of the relocations of radio-
marked animals were at the release site. In contrast, hard release
swift foxes tended to establish a den site away from the release
site. The distances, however, were not great. In 1987, radio-
marked foxes moved an average distance of 4.5 km before
establishing a den. One fox moved 50 km in 13 days before it was
shot. Arid conditions existed at the Alberta release site in 1988,
and only 4 of 12 radio-marked foxes remained on the study area.
These animals moved an average of 8 km before establishing a den
site. Herrero et al. (1990) concluded that under favorable
climatic conditions with an ample prey base, translocated swift
foxes tend to remain in the vicinity of the release site for both
soft and hard releases.

In the Alberta release, the mean number of days that radio-
marked foxes survived was 273. Thirty-seven foxes survived more
than 2 00 days. Predation was the principal cause of mortality
(63%) among recovered swift foxes and coyotes accounted for 63% of
the known predation. Golden eagles, bobcats, and undetermined
predators accounted for the remainder of the predation (Herrero et
al. 1990).

The swift fox is not yet considered established in Canada
despite known reproduction, survival of the young, and documented
breeding by these wild born foxes (Carbyn and Killaby 1989, Herrero
et al. 1990) . The Canadian reintroduction program has demonstrated
that swift foxes can be successfully released into former range
under favorable climatic conditions and with an adequate prey base.
The drought of 1988 resulted in dispersal of swift foxes away from
the release sites, and increased mortality. This was attributed to
a declining prey base. As of yet, the release program has not been
tested with a severe winter.

Both Carbyn and Killaby (1989) and Herrero et al. (1990)
caution that the prairie biome has been significantly altered and
that habitat conditions may no longer favor swift fox survival.
The loss of the bison (Bison bison) and wolf from the North
American prairie may have been a contributing factor in the loss of
the swift fox from its northern range. Bison may have been a
constant source of carrion for swift foxes, and wolves selectively
preyed upon coyotes keeping their numbers below present day levels.
Carbyn (1986) and Herrero et al. (1990) speculate that the swift
fox scavenged food at wolf kill sites, much as the arctic fox
( Alopex laqopus) does at polar bear (Urus maratimus) kill sites.

Both the South Dakota and Canadian reintroduction efforts
involved a pre-reintroduction survey of potential release sites.
This consisted of two Master level theses on the part of the
Canadians (Carlington 1980, and Reynolds 1983) . Sharps and
Whitcher (1984) list eight criteria for reintroduction sites used

35

in their program. These are: 1) available suitable habitat; 2) an
adequate food source, preferably prairie dogs; 3) a permanent water
source; 4) the absence of a predator poisoning program on, and
adjacent to, the ranch chosen as a release site; 5) restriction of
any furbearer or predator trapping; 6) release sites a minimum
distance of 24.0 km from major roads; 7) hunters having access to
the reintroduction area be made aware of the swift fox and be
instructed not to shoot them; and 8) coyote or red fox dens in the
immediate area would have to be eliminated prior to reintroduction.

Egoscue's (pers. comm.) recommendations concerning a
reintroduction were to release swift foxes in the vicinity of
recent sightings, release animals only when rabbits are abundant,
and control coyotes for several years until the swift fox are
established. Research conducted in South Dakota (Hillman and
Sharps 1978, Uresk and Sharps 1986, Sharps 1977, Sharps 1989)
indicates that black-tailed prairie dogs can serve as a constant
prey base. Swift foxes having access to prairie dogs would not
face the cyclic phenomenon of a lagomorph prey base, and would not
be limited by the lack of carrion as speculated by Herrero et al.
(1990) . Areas with extensive prairie dog colonies may serve as
indicators of suitable habitat for swift foxes.

Sharps (1977) estimated the operating expense of his 5 year
swift fox reintroduction/ecology program at $30,000. Twenty-five
percent was provided by the state of South Dakota and 75% by the
Federal government. The budget for a similar 5 year swift fox
reintroduction/ecology program in Alberta, Canada for the release
of 25 foxes was estimated at $140,000. To date, over $500,000 have
been spent on swift fox reintroduction in Canada.

Swift fox reintroduction is not an inexpensive program. The
South Dakota and Canadian experience shows that costs per released
animal may exceed $5,000, require that the program function for up
to a decade, and that survival of the swift foxes is low despite
the best efforts to assure success.

POTENTIAL MONTANA REINTRODUCTION SITES

Swift fox reintroduction, as evidenced by the Canadian
program, may be a long and expensive project with no guarantee of
establishing a population. The Montana prairie environment is much
changed from pre-historic times in which the swift fox evolved.
Species composition and numbers of the original ungulate population
is very different and both domestic and wild ungulates are now
closely managed with only minimal opportunity for carrion. Large
areas of prairie have been converted to small grain agriculture.
The gray wolf is gone and the coyote appears to have partly filled
its niche. The red fox is now abundant in what was formerly swift
fox range. Thus, the prey base, the habitat, and competition are

36

all much different now than a little more than 100 years ago. All
these factors may influence a reintroduction program.

A swift fox reintroduction into Montana is dependent upon
determining potential swift fox habitat. Although the original
distribution of the swift fox in Montana may never be known, it can
be surmised that swift foxes occurred in those regions of the state
where short and mid-grass prairies existed east of the continental
divide. Historical notes on swift fox distribution in Montana, and
its present range distribution would indicate that prairie country
with dense sagebrush or irregular topography may be marginal
habitat for swift fox. In addition, areas with extensive small
grain agriculture tend to support relatively few swift foxes as
opposed to native habitat in range areas where the swift fox is
still present (Fitzgerald et al. 1983, Stromberg pers. comm.).
Thus, agricultural areas may be eliminated as potential
reintroduction sites.

Habitat preference of the swift fox may overlap with other
prairie wildlife species. These more resilient species which have
persisted in Montana may serve as markers of what might be
potential swift fox habitat. Rabbits and hares (Sylvilaqus sp. ,
Lepus townsendii . and L. calif ornicus) figure significantly in
swift fox food habit studies and their presence would be a
prerequisite prior to any reintroduction. However, these species
may be more widely distributed in habitat preference than the swift
fox and could potentially occur in habitats unsuitable for the
swift fox.

Further refinement of potential swift fox habitat may be
judged by additional associated wildlife species. Fitzgerald et
al. (1983) reported the swift fox in northeastern Colorado to use
areas frequented by mountain plovers ( Charadrius montanus) .
Mountain plovers select those habitats with level topography and
the shortest of short grasses (Knowles et al. 1982) . Similarly,
swift foxes are documented to feed on black-tailed prairie dogs and
den in their colonies (Hillman and Sharps 1978) . In general,
prairie dogs select for areas with little topographic relief, short
grasses, and a sparse shrub layer. Areas with extensive complexes
of black-tailed prairie dog colonies (Clark et al. 1987) may be
good indicators of potential swift fox habitat.

We know that the swift fox historically existed in the Great
Falls - Fort Benton area, and the Blackfoot Indian Reservation.
The former area is primarily private land and has been extensively
converted to small grain agriculture. The modification of the
habitat makes this area undesirable for a swift fox reintroduction.
The land ownership pattern also would not be desirable for a
reintroduction. The habitat in the latter area is still largely
intact and appears to be suitable for a reintroduction. However,
the Reservation would present seemingly unique jurisdictional

37

situations which would add considerably to the complexity of a
reintroduction program.

Several other areas within the State may present favorable
situations for a swift fox reintroduction and should be considered
further. These are: 1) The UL Bend National Wildlife Refuge
represents an area of extensive Federal land ownership with no
legal trapping on Refuge lands. The habitat in this area appears
suitable with the prey base including prairie dogs and rabbits (For
additional information on this potential release site see FaunaWest
Wildlife Consultants (1986) and U.S. Fish and Wildlife Service
(1988).). 2) Areas of recent swift fox sightings (south of Miles
City, east of Glendive, and north of Broadus) should be evaluated
as possible reintroduction sites. The swift foxes captured at
these locations may have come from existing, nearby populations in
South Dakota and Wyoming. 3) Although the habitat in the area
immediately south of the Alberta/ Saskachtewan reintroduction site
is largely disturbed by agriculture, the presence of swift foxes
immediately north of Montana and the recent observations of swift
foxes south of the border suggests that this area should be
evaluated for a reintroduction.

A considerable amount of work has been done on laying the
ground work for a black-footed ferret reintroduction in Montana.
Much of the data collected from this work would be useful baseline
information for a swift fox reintroduction program. Moreover, the
concepts being developed for the black-footed ferret reintroduction
may be directly applicable to a swift fox reintroduction. In all
likelihood, the two reintroduction programs would be so similar
that they could be managed as sibling programs even though the
reintroductions may be temporally and spatially separated.

RECOMMENDATIONS

Two existing swift fox populations (one in South Dakota and
one in Wyoming) lie very close to Montana. A third population is
in the process of being established immediately north of Montana in
Alberta/ Saskatchewan. Over the past 12 years, at least four swift
foxes have been trapped in Montana. These four records are
spatially close to the existing out-of-state populations and could
represent dispersing animals or small resident populations. There
have also been several unconfirmed sightings which correspond
closely to the trapping records. This information provides a basis
from which to initiate a Montana swift fox investigation.

It is recommended that before any reintroduction program is
started, an effort be made to conduct an intensive survey for swift
foxes. This survey effort should be concentrated in those areas of
recent swift fox records and sightings. It is possible that one or
more existing small populations are already present in the State.

38

Jerry Choate (pers. comm.), based on his experience with swift
foxes and work in Montana, believes that it is highly likely that
a swift fox population is already established in Montana south and
east of the Yellowstone River. Three prime survey areas fitting
Choates • description and corresponding to recent records would be
east of Glendive, north of Broadus, and south of Miles City. A
fourth area warranting a swift fox survey would be north of Havre
adjacent to the area of the Canadian releases.

It is recommended that a survey for swift foxes consist of a
modified scent post survey (Linhart and Knowlton 1975) . The survey
should consist of 50 scent stations checked for 5 nights. Rather
than spacing the scent stations at 0.5 km intervals over a 24 km
course, they should be selectively placed along gravel and dirt
roads at about 1.6 km intervals. The scent stations would only be
placed in habitats that appear to be high potential for swift
foxes. The actual route for this survey would be established only
after making a cursory search of the survey area making note of
habitat, roads, and feasibility of the road for inclusion into the
route. One survey would be conducted in each of the four
geographic regions cited above as potentially containing swift
foxes. Late summer-early fall would be the best time period to
conduct this survey, both from a biological and meterological
basis. Additional survey work could be conducted concurrent with
the scent post survey. This should include interviews with
residents of the area, and night-time spotlight surveys along the
scent post survey routes.

Prior to conducting scent post surveys for swift fox presence
or absence data, Montana Dept. of Fish Wildlife and Parks should
consult with their own biologists, Bureau of Land Management, and
other interested parties on potential reintroduction sites.
Criteria established by Sharps and Whitcher (1984) should be used
as a guide for identifying potential reintroduction sites.
Available suitable habitat should be considered as two or more
townships of prairie habitat with less than 10% of the area
disturbed by agriculture. Landform should be level to gently
rolling. The area should be dominated by short grasses with only
a sparse shrub layer. Habitat suitability for swift foxes at
candidate release sites could be evaluated at the same time the
scent post surveys are run.

Legalizing the trapping harvest of swift fox in Montana for
one or more years would be an inexpensive means of gaining
presence/absence data on the swift fox in Montana. With a legal
harvest season, fur dealers would be more likely to report swift
foxes and incidents such as happened in 1984 (Henckel 1984) would
be avoided. With or without the legalization of trapping for swift
foxes, fur dealers within the potential range of the swift fox
should be guestioned annually for the next five years about swift
fox furs being bought or sold. Local, state, and Federal animal
damage control personnel in this region should also be questioned

39

concerning their knowledge of swift foxes, and be advised of the
importance of reporting sightings and records.

It is recommended that the Dept. of Fish Wildlife and Parks
establish a swift fox reporting system similar to that instigated
for the gray wolf (Flath 1979, USFWS 1980, Day 1981). Collection
and tabulation of these data over a period of years may result in
locating isolated populations and document the range expansion of
swift foxes in Montana.

ACKNOWLEDGMENTS

This project was conducted under contract with Montana
Department Fish, Wildlife and Parks. Arnold Dood supervised the
project and provided valuable logistical support and critically
reviewed the manuscript. The following people graciously provided
information or comments concerning the swift fox: Mark Boyce,
Ludwig Carbyn, Jerry Choate, Harold Egoscue, Stephen Herrero, James
Fitzgerald, Mark Johnson, Delbert Kilgore, Rick Pallister, Robert
Seabloom, Jon Sharps, and Mark Stromberg. We thank the many
curators of those museums which responded to our inquiry concerning
Montana swift fox specimens.

40

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reptiles, birds, mammals. Inst. Ecol. Stud. Univ. N. Dakota.
Res. Rep. 24 (REAP 78-4) . 549 pp.

Seton, E.T. 1929. Lives of game animals. Vol. I, Part II.
Doubleday, Dor an and Co., Inc. New York. x + 64 0 pp.

Sharps, J.C. 1977. The northern swift fox in South Dakota
(Unpublished) . Presented at Furbearer Management Workshop,
Rapid City, SD.

. 1984. Northern swift fox investigations 1977-1981. P-R

project SE-2 Study Number IV. South Dakota Dept. of Fish,
Game, and Parks. 2 6 pp.

. 1989. Swift fox inventory. Final Rep. to Rocky Mountain

Forest and Range Experiment Sta. Contract No. 43-82FT-9-
0920. 21 pp.

and M. F. Whitcher. 1984. Swift fox reintroduction

techniques. South Dakota Dept. of Game Fish, and Parks.
Rapid City. 31 pp.

Snow, C. 1973. San Joaquin kit fox, Vulpes macrotis mutica.
Related subspecies and the swift fox, Vulpes velox. Habitat
management series for endangered species. Report No. 6. BLM
Tech. Note. Denver Service Center. 24 pp.

47

Soper, J.D. 1964. The mammals of Alberta. Hanly Press Ltd.
Edmonton. 402 pp.

Stiles, C.W. and C.E. Baker. 1935. Key-catalogue of parasites
reported for Carnivora with their possible health
importance. Bull. U.S. Nat. Inst. Health, 163:913-1223.

Stromberg, M.R. , and M.S. Boyce. 1986. Systematics and
conservation of the swift fox, Vulpes velox, in North
America. Biol. Conser. , 35:97-110.

Swanson, G.A. , T. Surber, and T.S. Roberts. 1945. The mammals of
Minnesota. Minnesota Dept. Conserv. Div. Game and Fish Tech.
Bull., 2:1-108.

Thompson, L.S. 1982. Distribution of Montana Amphibians,
Reptiles and Mammals. Montana Audubon Concil, Helena. 25
pp.

Thorne, E.T. and E. Williams. 1988. Diseases and endangered
species: the black-footed ferret as a recent example.
Conservation Biology, 2:66-73.

Thornton, W.A., and G.C. Creel. 1975. The taxonomic status of
kit foxes. Texas J. Sci., 26:127-136.

, , and R.E. Trimble. 1971. Hybridization in the fox

genus Vulpes in west Texas. Southwest. Nat., 15:473-484.

Uresk, D.W. and J.C. Sharps. 1986. Denning habitat and diet of
the swift fox in western South Dakota. Great Basin Nat.,
46:249-253.

U.S. Fish and Wildlife Service. 1979. Federal Register, 17 June
1979, p. 3638.

. 1980. Northern Rocky Mountain Wolf Recovery Plan.

. 1982. Federal Register, 30 December 1982. pp. 58454-60.

. 1988. Reintroduction of the swift fox (Vulpes velox) to

the U.L. Bend National Wildlife Refuge. Environmental
Assessment. 14 pp.

. 1989. List of threatened and endangered species. Federal

Register.

. 1990. Report on the status of the swift fox, Vulpes

velox. a Category 2 Taxon, in North Dakota. North Dakota
State Office, Bismarck. 18 pp.

48

Vallard, D. 1985. Rare fox trapped near Glendive. Miles City
Star, 20 January.

Van Ballenberghe, V. 1975. Recent records of the swift fox
(Vulpes velox) in South Dakota. J. Mamm. , 56:525.

Visher , S.S. 1914. The biology of Harding County, Northwestern
South Dakota. S.D. Geographic Survey, 6:126.

Voigt, D.R. and B. Earle. 1983. Avoidance of coyotes by red fox
families. J. Wildl . Manage., 47:852-857.

Weidman, F.D. 1915. Coccidium biqeminum stiles in swift foxes.
J. Comp. Pathol. Therap. , 28:320-323.

Yates, T.L., W.R. Barber, D.M. Armstrong. 1987. Survey of North
American collections of recent mammals. J. Mammal.,
Supplement to Vol. 68 No. 2. 7 6 pp.

Young, S.P. 1951. Part 1, Its history, life habits, economic

status, and control. Pages 1-226. in S.P. Young and H.H.T.

Jackson. The clever coyote. Wildl. Manage. Inst.,
Washington, D.C. 411 pp.

Zegers, D.A. 1976. Predator abundance surveys and the status of
the swift fox (Vulpes velox) in southeastern Colorado.
Unpubl. manuscript. 76 pp.

Zumbaugh, D.M. 1984. Natural history of foxes in Kansas. MS
Thesis. Fort Hays State Univ. Hays, Kansas.

, and J.R. Choate. 1985. Historical biogeography of foxes

in Kansas. Trans, of the Kansas Acad, of Sci . , 88:1-13.

, , and L.B. Fox. 1985. Winter food habits of the swift

fox on the central high plains. Prairie Nat., 17:41-47.

49

[

APPENDIX A

LIST OF MONTANA SWIFT FOX SPECIMENS HELD IN MUSEUMS

28 MAR 1990

SMITHSONIAN INSTITUTION

MAMMALS MASTER LIST

USNM Number: 108260

Fami ly Code: 6010

Family: CANIDAE

Genus: VULPES

Species: VELOX

Subspecies: VELOX

Date Col 1 : 24 MAR 1901

1st Geo Div: UNITED STATES

2nd Geo Div: MONTANA

3rd Geo Div: TETON COUNTY

Specific Loc: BLACKF00T

Loc Modifier: KIPP POST OFFICE

Collector: DAWSON, J.

Field Number: NO NUMBER

Remarks: 2905 X

Sex : 6

Preparation: SKIN AND SKULL

USNM Number: 108318

Fami ly Code: 6010

Family: CANIDAE

Genus: VULPES

Species: VELOX

Subspecies: VELOX

Date Col 1 : 08 APR 1901

1st Geo Div: UNITED STATES

2nd Geo Div: MONTANA

3rd Geo Div: TETON COUNTY

Specific Loc: BLACKFOOT

Loc Modifier: KIPP POST OFFICE

Collector: DAWSON, J.

Field Number: NO NUMBER

Remarks: 2909 X

Sex: $

Preparation: SKIN AND SKULL

USNM Number: 108319

Fami ly Code: 6010

Family: CANIDAE

Genus: VULPES

Species: VELOX

Subspecies: VELOX

Date Col 1 : 28 MAR 1901

1st Geo Div: UNITED STATES

2nd Geo Div: MONTANA

3rd Geo Div: TETON COUNTY

Specific Loc: BLACKFOOT

Collector: MONROE, J. B.

Field Number: NO NUMBER

Remarks: SKIN GIVEN AS INDEFINITE LOAN
CONTROL SUPERVISOR, AMHERST,

Sex: ?

Preparation: SKULL ONLY

USNM Number : 1 1 6560

Locator: FRV

Bldg / Area: FUR VAULT

Fami ly Code: 6010

Family: CANIDAE

Genus: VULPES

Species: VELOX

Subspecies: VELOX

Date Coll : -- MAR 1901

1st Geo Div: UNITED STATES

2nd Geo Div: MONTANA

3rd Geo Div: TETON COUNTY

Specific Loc: BLACKFOOT

Loc Modifier: KIPP POST OFFICE

Collector: DAWSON, J.

Field Number: NO NUMBER

Remarks: 3502 X

Sex: ?

Preparation: SKIN ONLY

USNM Number: 1 16561

Locator: FRV

Bldg / Area: FUR VAULT

Fami ly Code: 6010

Family: CANIDAE

Genus: VULPES

Species: VELOX

Subspecies: VELOX

TO MAMMAL

Date Col 1 :

-- MAR 1901

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

1 Mr\r\ i f i *
LUt I'lUU 1 1 I CI •

KIPP Pfl^T nFFITF

Col lector :

DAWSON, J.

r ie la Number :

NU NUMBhK

Remarks :

3503 X

Sex :

Preparation:

SKIN AND SKULL

USNM Number :

1 1*587^

Fami ly Code:

6010

Fami ly:

CANIDAE

Genus :

VULPES

Species :

VELOX

Subspecies :

VELOX

Date Col 1 :

20 NOV 1905

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Spec i f i c Loc :

D L AUrVr UU 1

Col lector :

MONROE, J. B.

Field Number :

NO NUMBER

Remarks :

5690 X

Sex :

9

Preparation:

SKIN AND SKULL

USNM Number:

H»5875

Fami ly Code:

6010

Fami ly:

CANIDAE

Genus :

VULPES

Spec i es :

VELOX

Subspecies :

VELOX

Date Col 1 :

20 DEC 1905

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

DLACKrUU 1

Col lector :

MUNROE, G. J.

Field Number :

NO NUMBER

Remarks :

5619 X

Sex :

9

Preparation:

SKIN AND SKULL

USNM Number:

1^5877

Fami ly Code:

6010

Fami ly:

CANIDAE

Genus :

VULPES

Species :

VELOX

Subspecies :

VELOX

Date Col 1 :

23 DEC 1905

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

u L Mur\ r UU 1

Col lector :

MUNROE, d. K.

Field Number :

l i r-\ nip r~ r"i

NO NUMBER

Remarks :

5693 X

Sex :

c*

Preparation:

SKIN AND SKULL

USNM Number:

14631 1

Fami ly Code:

6010

Fami ly:

CANIDAE

Genus :

VULPES

Species :

VELOX

Subspecies :

VELOX

Date Coll :

17 DEC 1905

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

Col lector :

MONROE , J. B.

Field Number:

1

Remarks :

ETC 1 y
O f D 1 A

Sex :

d

Preparation:

SKIN AND SKULL

USNM Number:

11*6312

Fami ly Code:

6010

28 MAR 1990

SMITHSONIAN INSTITUTION

MAMMALS MASTER LIST

F sm i 1 y :

CANIDAE

Genus :

VULPES

Spec i es :

VELOX

Subspec i es *

VELOX

Date Col 1 :

28 DEC 1905

1 «; t Gf»o D i v

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKF00T

Col lector :

MONROE, J. B.

Field Number:

2

Remarks :

5762

Sex :

d

Preparation:

SKIN AND SKULL

146313

Fami ly Code:

fin 1 n

OU IU

F ami 1 y :

r A M t n A F

Genus :

VIII PFS

V UL r C O

Spec i es :

V C L UA

Subspecies :

)/fi nx

V L L UA

Ua ie v-O 1 1 •

1 S> I UcO U 1 V .

IIMTTFn ^TATF^

' 1 . J. 1 Li1 J 1 M 1 CO

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Speci f ic Loc :

BLACKFOOT

Col lector :

MONROE, J. B.

Field Number :

161

Remarks :

5766 X

Sex :

6*

Preparation:

SKIN AND SKULL

t I^NM Ni imh^r *

146315

F s> m i 1 \ / i i~\t~iQ '

r am i i y tuuc .

fin 1 n

F am i 1 y ■

("ANIDAE

VULPES

VELOX

VELOX

Dat^ fnl 1 ■

Ua IC \s \J i l •

Ofi JAN 1Q06

let Ron H i \j '

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

Col lector :

MONROE, J. B.

Field Number :

1 1

Remarks :

5771 X

Sex :

d

Preparat i on :

SKIN AND SKULL

146316

r tain i i y wtjutr .

60 1 0

TANTDAF

F Genus '

VUL PES

VELOX

Q t i o i""iA^"" i a c ■
OUUofJCt 1 C9 •

VELOX

1fl ,IAN 1Q0fi

let fioA Hi u ■

IINTTFfi ^TATF9

UlvX 1 1. U j 1 W 1 CO

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Spec i f i c Loc :

BLACKFOOT

Co 1 lector :

MONROE, d. B.

Field Number:

14

Remarks :

5774 X

Sex :

a

Preparat i on :

SKIN AND SKULL

146317

60 1 0

F ami 1 y :

TAN1 nAF

Genus :

Will PFS

opec l ca •

vfi nx

ua ic vU i i .

m nFf 1 QHR

let fior> Div

1 I UiU U 1 v .

unttfd <;tatf<;

Uli 1 1 LU O 1 M 1 CO

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

Col lector :

MONROE, J. B.

Field Number :

3

Remarks :

5763 X

Sex :

9

Preparation:

SKIN AND SKULL

USNM Number:

146318

Fami ly Code:

6010

Fami ly:

CANIDAE

Genus :

VULPES

Species :

VELOX

Subspecies :

VELOX

Date Col 1 :

30 DEC 1905

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

Col lector :

MONROE, d. B.

Field Number :

4

Remarks :

5764 X

Sex :

Preparation:

SKIN AND SKULL

USNM Number :

146319

F ami 1 y Code :

6010

Fami ly :

CANIDAE

Genus :

VULPES

Spec i es :

VELOX

Subspecies :

VELOX

Date Col 1 :

31 DEC 1905

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

Collector:

MONROE, d. B.

Field Number :

5

Remarks :

5765 X SKULL MISSING

Sex :

9

Preparat i on :

SKIN AND SKULL

USNM Number :

146320

Fami ly Code :

6010

Fami ly:

CANIDAE

Genus :

VULPES

Species :

VELOX

Subspeci es :

VELOX

Date Col 1 :

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

Col lector :

MONROE, G. G.

Field Number :

7

Remarks :

5767 X SKULL MISSING

Sex :

9

Preparation:

SKIN AND SKULL

USNM Number :

146321

Fami ly Code :

6010

Fami ly:

CANIDAE

Genus :

VULPES

Species :

VELOX

Subspecies :

VELOX

Date Col 1 :

23 DEC 1905

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

Col lector :

MONROE , d. B.

Field Number:

9

Remarks :

5769 X

Sex:

9

Preparation:

SKIN AND SKULL

USNM Number :

146322

Fami 1 y Code :

6010

Fami ly:

CANIDAE

Genus :

VULPES

Spec i es :

VELOX

Subspecies :

VELOX

Date Col 1 :

07 JAN 1906

1st Geo Div:

UNITED STATES

2nd Geo Div:

MONTANA

3rd Geo Div:

TETON COUNTY

Specific Loc:

BLACKFOOT

Col lector :

MONROE , d. B.

Field Number :

10

zb mak iyyu

SMI IHSONIAN INSTITUTION
MAMMALS MASTER LIST

Remarks '.

D / / U A

P i e> 1 rl Mi imhar *

r Ic IU iMUiiiLrtrr

oex .

o

Doma r U c ■
nClllol r\© .

4634 X

Prepar a t i on

C.K TM AND C.KI II 1

Sex :

0

+

Pr «r"\5i r* si t i nn •
rl ClJot a L 1 \Ji 1 .

C.K T N AND C. K 1 J I 1

USNM Number:

l 40323

133093

Fami 1y Code :

6010

USNM Number:

Fami ly:

CANIDAE

Fami ly Code:

6010

Genus :

VULPES

Fami ly:

CANIDAE

Species :

VELOX

Genus :

VULPES

Subspecies :

VELOX

Species :

VELOX

Date Col 1 :

14 JAN 1906

Subspecies :

VELOX

1st Geo Div:

UNITED STATES

Date Col 1 :

20 dUN 1904

2nd Geo Div:

MONTANA

1st Geo Div:

UNITED STATES

3rd Geo Div:

TETON COUNTY

2nd Geo Div:

MONTANA

Specific Loc:

BLACKFOOT

3rd Geo Div:

TETON COUNTY

Col lector :

MONROE, d. B.

Specific Loc:

KIPP

Field Number :

12

Col lector :

MONROE , J . B .

Remarks :

5772 X

Field Number :

3

Sex :

9

Remarks :

4635 X

Preparation:

SK I N AND SKULL

C w •

oex :

9

Prepara t i on ;

iMPJ ANU oRULL

USNM Number:

A0**2 1 3

133094

Fami ly Code:

6010

USNM Number:

Fami ly:

CANIDAE

Fami ly Code:

6010

Genus :

VULPES

Fami ly:

CANIDAE

Species :

VELOX

Genus :

VULPES

Subspecies :

VELOX

Species :

VELOX

Date Col 1 :

Subspecies :

VELOX

1st Geo Div:

UNITED STATES

Date Coll:

20 dUN 1904

2nd Geo Div:

MONTANA

1st Geo Div:

UNITED STATES

Specific Loc:

FORT BENTON

2nd Geo Div:

MONTANA

Lo 1 lecxor .

Kill 1 1 1 AD

MULLAK

o p o ueo u i v .

TCTnM rni imtv
1 1 1 UlN LUUlM 1 1

Field Number :

1056

Specific Loc:

KIPP

Sex :

Col lector :

ynnnnr 1 D

MONROE. , J . B .

Preparat ion:

SKULL ONLY

Field Number :

4

Remarks :

4636 X

USNM Number:

1 30059

Sex :

9

Fami ly Code:

6010

Preparation:

SKIN AND SKULL

Fami ly:

CANIDAE

135129

Genus :

VULPES

USNM Number:

Species :

VELOX

Fami ly Code:

6010

Subspecies :

VELOX

Fami ly:

CANIDAE

Date Col 1 :

20 OCT 1903

Genus :

VULPES

1st Geo Div:

UNITED STATES

Species :

VELOX

2nd Geo Div:

MONTANA

Subspecies :

VELOX

3rd Geo Div:

TETON COUNTY

Date Col 1 :

20 OCT 1904

Specific Loc:

KIPP

1st Geo Div:

UNITED STATES

Col lector :

MONROE, J. B.

2nd Geo Div:

MONTANA

Meld Number :

tin itii m prn

NO NUMBER

3rd Geo Div:

TETUN CUUNTY

Remarks :

4656 X

Specific Loc:

KIPP

C a w •

oex .

0
¥

Col lector :

MUNKUt , u - D .

Prepar a t i on :

Clf T M AKin Cfcf 1 II 1

SMN ANU SfVULL

Field Number:

Kin Ml III D C D

NO NOMbtK

Remarks :

USNM Number:

133091

Sex :

6

Fami ly Code:

6010

Prepar at i on :

C W T M A kin C fcf 1 1 1 1

SKIN AND 5KULL

Fami ly:

CANIDAE

13o5oo

Genus :

VULPES

USNM Number:

Species :

VELOX

Fami ly Code:

6010

Subspecies :

VELOX

Fami ly:

CANIDAE

Date Col 1 :

18 JUN 1904

Genus :

VUL PES

1st Geo Div:

UNITED STATES

Species :

VELOX

2nd Geo Div:

MONTANA

Subspecies :

VELOX

3rd Geo Div:

TETON COUNTY

Date Coll:

17 MAR 1905

Specific Loc:

KIPP

1st Geo Div:

UNITED STATES

Col lector :

MONROE, d. B.

2nd Geo Div:

MONTANA

Field Number :

NO NUMBER

3rd Geo Div:

TETON COUNTY

Remarks :

4633 X

Specific Loc:

KIPP

Sex :

?

Col lector :

uniinnr 1 b

MONROE , d . B .

Prepar a t i on :

dMN ANU SKULL

Field Number :

NO NUMBER

Remarks :

5 1 92 X

USNM Number:

133092

Sex :

ri
¥

Fami ly Code:

6010

Prepar at i on :

C W T M A nr\ C IS I II 1

SKIN AND SKULL

Fami ly :

CANIDAE

139185

Genus :

VULPES

USNM Number:

Species :

VELOX

Fami ly Code:

6010

Subspecies :

VELOX

Fami ly:

CANIDAE

Date Col 1 :

19 JUN 1904

Genus :

VULPES

1st Geo Div:

UNITED STATES

Species :

VELOX

2nd Geo Div:

MONTANA

Subspecies :

VELOX

3rd Geo Div:

TETON COUNTY

Date Col 1 :

01 AUG 1905

Spec i f i c Loc :

KIPP

1st Geo Div:

UNITED STATES

Col lector :

MONROE, d. B.

2nd Geo Div:

MONTANA

28 MAR 1990

SMITHSONIAN INSTITUTION
MAMMALS MASTER LIST

3rd Geo Div:

TETON COUNTY

1 s t Geo Div:

UNITED STATES

Specific Loc :

KIPP

2nd Geo Div:

MONTANA

r*n 1 lor1 tor '

MONROE J B

?rri fi^o Div

J 1 w UCv L/ P V .

TETON COUNTY

Field Number :

NO NUMBER

Specific Loc:

KIPP

Remarks :

5612 X

Col lector :

MUNROE, J. K.

Sex :

$

Field Number :

NO NUMBER

Prepar a t i on :

SKIN ONLY

Remarks :

5682 X

Sex :

6

USNM Number :

139 186

Preparation:

SKIN AND SKULL

Fami ly Code:

6010

Fami ly:

CAN I DAE

USNM Number :

Genus :

VULPES

Fami 1 y Code :

60 10

Species :

VELOX

Fami ly:

CANIDAE

Subspecies :

VELOX

Genus :

VULPES

Date Col 1 :

27 AUG 1905

Species :

VELOX

1st Geo Div:

UNITED STATES

Subspecies :

VELOX

2nd Geo Div:

linilT A IlI a

MONTANA

Date Col 1 :

ut ii 4 ft r\c

-- WIN 1905

3rd Geo Div:

TETON COUNTY

1st Geo Div:

UNITED STATES

Specific Loc:

KIPP

2nd Geo Div:

MONTANA

^rrl Cif*r\ Div"

Jl \J UCU L* 1 V .

TETON COUNTY

Field Number:

NO NUMBER

Specific Loc:

KIPP

Remarks :

5613 X

Co 1 1 ec tor :

MUNROE , d . K .

Sex :

c5

Field Number:

NO NUMBER

Prepar a t i on :

SKIN ONLY

Remarks :

5559 X

Sex :

?

USNM Number :

1 3Q1R7

Prepar a t i on :

SKIN AND SKULL

Fami ly Code:

6010

Fami ly:

CANIDAE

MCilltl 1. 1 1

USNM Number :

Genus :

VUL PES

Fami ly Code:

60 10

Species :

VELOX

Fami ly:

P1 A 11 T Pi A r

CANIDAE

Subspecies :

VELOX

Genus :

VUL PES

Date Col 1 :

r\ r\ All/*1 4 flftC

03 AUG 1905

Species :

VELOX

1 s t Geo Div:

UNITED STATES

Subspecies :

VELOX

2nd Geo Div:

uniiT A LIA

MONTANA

Date Col 1 :

1905

3rd Geo Div:

TETON COUNTY

1st Geo Div:

1 ItlTTTPt PTATPf

UNITED STATES

Specific Loc:

KIPP

2nd Geo Div:

MONTANA

MDNRnF il R

J I U UCU V 1 V *

UPPER TETON COUNTY

Field Number :

NO NUMBER

Specific Loc:

KIPP

Remarks :

5615 X

Co 1 1 ec tor :

LANTZ, D. E.

Sex :

Field Number:

NO NUMBER

Prepar a t i on :

SKIN ONLY

Remarks :

5784 X

Sex :

?

USNM Number :

Prepar a t i on :

SKULL ONLY

Fami ly Code:

6010

Fami ly:

CANIDAE

USNM Number :

Genus :

VULPES

Fami ly Code:

6010

Species :

VELOX

Fami ly :

CANIDAE

Subspecies :

VELOX

Genus :

VULPES

Date Col 1 :

22 DEC 1905

Species :

VELOX

1 s t Geo Div:

IIHTTPn PTATPf

UNITED STATES

Subspecies :

VELOX

2nd Geo Dw:

AirM.IT A k 1 A

MONTANA

Date Col 1 :

MAY 1905

JiO UCU V 1 V .

TFTnw rnnwTY

IIMTTFn 'nTATF's

Spec i f i c Loc :

KIPP

2nd Geo Div:

MONTANA

MIINRflF M

Jl U UCU L' 1 V ,

TETON COUNTY

Field Number :

NO NUMBER

Specific Loc:

KIPP, [UPPER TETON

Sex :

Co 1 lector :

MONROE , J . B .

Prepar a t i on :

SKIN AND SKULL

Field Number :

NO NUMBER

Remarks :

5556 X

USNM Number :

1 1»622&

Sex :

Fami ly Code:

<5U 1 0

Prepara t i on :

SKIN ONLY

Fami ly:

CANIDAE

1 tOJUJ

Genus :

VUL PES

■ ■ #» Lift* A. 1 L_ -

USNM Number :

Species :

VELOX

Fami ly Code:

60 10

Subspecies:

VELOX

Fami ly:

CANIDAE

Date Col 1 :

A A llflW 4 A AC

04 NOV 1905

Genus :

VULPES

1st Geo Div:

1 IMTTFPi f TATTf

UNITED STATES

Species :

VELOX

2nd Geo Div:

MONTANA

Subspecies :

VELOX

3rd Geo Div:

TETON COUNTY

Date Col 1 :

1905

OpCClIlC LOC •

KIPP

i s x beo LM V .

IIMTTCn CTATCC
UNI 1 tV j 1 A 1 to

Co 1 1 &c tor ■

MUNROE , d. K.

4ml IU Uvv U } V •

MONTANA

fslfl NIIMRFR

1 'i U IiUI<IuLi>

o r u oeo uiVi

TETON COUNTY

Remarks :

5681 X

Spec i f i c Loc :

KIPP, [UPPER TETON

Sex :

6

Co 1 lector :

MONROE, d. B.

Prepar at i on :

SKIN AND SKULL

Field Number:

NO NUMBER

Remarks :

5783 X

USNM Number:

146227

Sex :

?

Fami ly Code:

6010

Preparation:

SKULL ONLY

Fami ly:

CANIDAE

1W97

Genus :

VULPES

USNM Number:

Species :

VELOX

Fami ly Code:

6010

Subspecies :

VELOX

Fami ly:

CANIDAE

Date Coll:

07 NOV 1905

Genus :

VULPES

W I w> v> U

dim mOUIUHIM 1IM3I1IUI1UN

MAMMALS MASTER LIST

Species
Subspecies
Date Coll
1st Geo Div
2nd Geo Div
3rd Geo Div
Specific Loc
Col lector
Field Number
Remarks
Sex

Preparation

USNM Number
Fami ly Code
Fami ly
Genus
Species
Subspecies
Date Col 1
1st Geo Div
2nd Geo Div
3rd Geo Div
Specific Loc
Col lector
Field Number
Remarks
Sex

Preparation

USNM Number
Fami ly Code
Fami ly
Genus :
Spec i es :
Subspecies :
Date Col 1 :
1st Geo Div:
2nd Geo Div:
3rd Geo Div:
Specific Loc:
Col lector :
Field Number :
Remarks :
Sex :

Preparation:

USNM Number:
Fami ly Code:
Fami ly:
Genus :
Species :
Subspecies :
Date Col 1 :
1st Geo Div:
2nd Geo Div:
3rd Geo Div:
Specific Loc:
Col lector :
Field Number :
Remarks :
Sex :

Preparation:

USNM Number:
Fami ly Code:
Fami ly:
Genus :
Spec i es :
Subspecies :
Date Col 1 :
1st Geo Div:
2nd Geo Div:
3rd Geo Div:
Specific Loc:
Col lector :
Field Number :
Remarks :
Sex :

Preparation:

VELOX
VELOX

-- MAY 1905
UNITED STATES
MONTANA
TETON COUNTY

KIPP, [UPPER TETON COUNTY]
MONROE, d. B.
NO NUMBER

5556 X

?

SKIN ONLY

147767

6010

CANIDAE

VULPES

VELOX

VELOX

1904

UNITED STATES
MONTANA
TETON COUNTY

KIPP, (UPPER TETON COUNTY]
MONROE. d. B.
NO NUMBER
5558 X

?

SKIN AND SKULL

146324

6010

CANIDAE

VULPES

VELOX

VELOX

20 DEC 1905

UNITED STATES

MONTANA

TETON COUNTY

UPPER TETON COUNTY

MONROE, d . B.

13

5773 X

9

SKIN AND SKULL

146325

6010

CANIDAE

VULPES

VELOX

VELOX

25 JAN 1906

UNITED STATES

MONTANA

TETON COUNTY

UPPER TETON COUNTY

MONROE, d. B.

15

5775 X

9

SKIN AND SKULL

146372

6010

CANIDAE

VULPES

VELOX

VELOX

28 dUN 1905

UNITED STATES

MONTANA

TETON COUNTY

UPPER TETON COUNTY

MONROE, d. B.

1

5555 X

9

SKIN ONLY