Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

Suggested Citation:

Harrell, D., and L. Marks. 2009. Habitat selection and changes in the white-tailed 

and black-tailed prairie dog population within the northern Bighorn Basin, 

Wyoming, Technical Note 431. U.S. Department of the Interior. Bureau of Land 

Management, Cody Field Office, Wyoming. BLM/WY/ ST-09/031+1110. 16 pp.

Production services provided by:

Bureau of Land Management

National Operations Center

Information and Publishing Services Section

P.O. Box 25047

Denver, Colorado 80225

Available online at: 

www.blm.gov/wy/st/en/field_offices/Cody.html

BLM/WY/ ST-09/031+1110

Cover photo (white-tailed prairie dog) and all other photos except as noted by:

Destin Harrell, BLM Cody Field Office
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

By:

Destin Harrell

Wildlife Biologist

Bureau of Land Management

Cody Field Office, Wyoming

Lisa Marks

Geologist

Bureau of Land Management

Cody Field Office, Wyoming

White-tailed prairie dog
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

iii

WE WOULD LIKE TO THANK Joe Capodice, 

retired biologist for the BLM. He helped 

inspire our prairie dog survey. Steve Kiracofe 

was a big help for the soil survey analysis as 

was Richard Lowry with statistical questions. 

Field crew members who were kind enough 

to collect much of the data for this publication 

include Brian Latta, Bill Palmeroy, Katie 

VinZant, and Darci Stafford. Dean Biggins, 

Dennis Saville, Bob Oakleaf, and Martin 

Grenier offered much helpfuladviceand 

review. Bill Wilson was always there for 

GPS and GIS data processing. Mike Stewart 

was generous in offering reviews, ideas, 

and support. Kathy Rohling and Janine 

Koselak at the National Operations Center 

contributed invaluable efforts by editing, 

arranging, and drawing original artwork for 

this paper. And finally, thanks to the Bureau 

of Land Management’s Cody Field Office 

staff and management for helping with this 

collaborative effort.

ACKNOWLEDGMENTS ACKNOWLEDGMENTS

Black-tailed prairie dog, Randy Hayes, BLM
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

v

ABSTRACT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . vii

INTRODUCTION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .1

STUDY AREA . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3

METHODS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5

RESULTS . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .7

Population Demographics . . . . . . . . . . . . . . . . . . . . .7

Habitat Associations . . . . . . . . . . . . . . . . . . . . . . . . . .9

DISCUSSION . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .11

LITERATURE CITED . . . . . . . . . . . . . . . . . . . . . . . . . . . . .15

CONTENTS CONTENTS

Burrowing owls
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

vi

Burrowing owls, a prairie dog associate, frequently nest in empty burrows.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

vii

WITHIN THE NORTHERN BIGHORN BASIN, 

Wyoming, prairie dog (Cynomys) survey 

data collected between 1980 and 1989 were 

compared with data from a 2001–2005 

survey. From these data, distribution, 

activity level, and habitat selection based 

on range site and precipitation categories 

were analyzed. Overall, the white-tailed 

prairie dog (C. leucurus) population declined 

in distribution and activity. Occupied active 

area decreased 71 percent from 7,031.8 ha 

(17,374.9 ac) to 2,011.5 ha (4,970.6 ac); 

there was also a 37 percent decrease in 

active town abundance and a 55 percent 

decrease in the mean active town area. 

Declines in occupied area for the Meeteetse 

Metapopulation (87 percent) were greater 

than declines for the remaining population 

(Basin Metapopulation - 39 percent). There 

was also a very small population of blacktailed prairie dogs (C. ludovicianus) that 

increased 8 percent to 77.4 ha (191.3 ac). 

Density of active burrows declined 

significantly (P < 0.006) for the Basin 

Metapopulation. The mean nearest 

neighbor distances separating white-tailed 

prairie dog towns for both surveys were 

approximately 2.0 km. Towns appear to be 

significantly associated with saline upland, 

loamy, sandy, shallow loamy, and shale 

range sites (P < 0.0001). White-tailed prairie 

dogs were also significantly associated 

with precipitation zones (P < 0.0001) and 

may prefer the 20.3–25.4 cm (8–10 in) 

precipitation zone even though they 

were more common in the 12.7–22.9 cm 

(5–9 in) precipitation zone. Other variables 

not measured also likely contribute to the 

observed distribution of white-tailed prairie 

dog town site selection.

ABSTRACT ABSTRACT

White-tailed prairie dog
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

Landscape of the Meeteetse prairie dog metapopulation
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

1

PRAIRIE DOGS (Cynomys spp.), are often 

described as a keystone species and 

ecological engineers (Miller et al. 1994; 

Bangert and Slobodchikoff 2000), have 

declined as much as 98 percent throughout 

North America since European settlement 

(Ceballos and Reading 1994). Viability of 

several species has become an important 

biodiversity issue. All species are listed or 

have been petitioned for listing under the 

Endangered Species Act of 1973.

This biodiversity issue is relevant in the 

Bighorn Basin, Wyoming, where whitetailed prairie dogs (C. leucurus) are near the 

northern extent of their range and where 

four black-tailed prairie dog (C. ludovicianus)

towns are grouped together near the city of 

Cody, Wyoming (Seglund et al. 2004). The 

origin of these sympatric black-tailed prairie 

dogs, isolated within the white-tailed 

prairie dog range is unknown. Speculation 

includes release from captivity or an isolated 

population of their former range.

White-tailed prairie dogs and black-tailed 

prairie dogs have different life histories and 

may influence the ecosystem in different 

ways. White-tailed prairie dogs may clip 

vegetation less than black-tailed prairie 

dogs (Clark 1973). During the winter, whitetailed prairie dogs may enter torpor at an 

ambient air temperature of 7° C for longer 

periods of time than black-tailed prairie 

dogs, which are facultative hibernators 

during winter (Harlow and Menkens 1986; 

Lehmer and Biggins 2005). Both species 

alter their environment, providing unique 

habitat for associated species, and can be a 

substantial prey base for predators (Campbell 

and Clark 1981; Knowles et al. 1982; Miller 

et al. 1994; Weltzin et al. 1997).

The black-footed ferret (Mustela nigripes)

preys upon prairie dogs for the majority of 

its diet and is one of many species associated 

with and adapted to prairie dog towns 

(Hillman et al. 1979; Campbell and Clark 

1981; Cambell et al. 1987). During the 

evening of September 26, 1981, a blackfooted ferret was killed by a rancher’s dog 

approximately 48 km (30 miles) south 

of Cody and a few kilometers west of 

Meeteetse, Wyoming; the consequential 

live capture of the once thought extinct 

black-footed ferret occurred in October 1981 

(Menkens and Anderson 1991). In 1985, 

sylvatic plague (Yersinia pestis) was 

discovered in the prairie dog metapopulation 

near Meeteetse where the population of 

black-footed ferrets was discovered 

(Menkens and Anderson 1991).

These discoveries helped to prompt James 

Bredy and Steven Coy of the Bureau of Land 

INTRODUCTION INTRODUCTION

Mountain plover, endemic to North America, find prairie dog towns to be good nesting habitat.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

2

Management’s (BLM) Cody Resource Area to 

begin surveys for prairie dogs. Surveys were 

conducted between 1980 and 1989 using 

a variety of techniques for locating and 

mapping white-tailed prairie dog colonies 

including, but not limited to, infrared aerial 

photography, helicopter flights, personal 

communications, and ground observations. 

These data were used to inventory prairie 

dog towns, better identify potential habitat 

for black-footed ferrets, and to produce a 

Prairie Dog Ecosystem Habitat Management 

Plan (Coy and Roberts 1985). Their study area 

excluded the Meeteetse Metapopulation, 

which was mapped independently by the 

U.S. Fish and Wildlife Service (FWS) on 

1:100,000 maps in 1988 (Coy and 

Roberts 1985).

This report compares data from the survey 

conducted by the BLM in 2001–2005 with 

the 1980–1989 surveys conducted by the 

BLM and FWS, and summarizes changes in 

indices of distribution and abundance. The 

2001–2005 data were also used for analysis 

of habitat associations. Methods were used 

similar to those in 1980–1989, which made 

comparisons possible with acknowledgement of biases, limitations, and errors when 

replicating historical surveys. These data 

may help identify trends in populations; 

however, these trends do not identify the 

many factors that may affect prairie dog 

distribution and abundance.

Prairie dog distribution and activity may be 

regulated by their natural history, which is 

different for white-tailed and black-tailed 

prairie dogs (Hoogland 1996). Prairie dog 

towns are not randomly distributed and 

towns may be clumped (Hillman et al. 

1979). Prairie dogs produce one litter per 

year with an average of 3.08 to 3.88 pups, 

and survival may be less than 60 percent in 

the first year (Clark and Stromberg 1987; 

Hoogland 2001). Female harassment may 

be caused by a shortage of unrelated females 

in the coterie, encouraging dispersal (Garrett 

and Franklin 1988). In Albany County, 

Wyoming, pups began to disperse at the 

time when the population density was 

greatest (late June and early July) (Clark 

1973). Young, dispersing males tended to 

occupy old, uninhabited burrow systems 

(Clark 1973). Some adults may also emigrate, 

leaving the young behind, possibly due to 

the animosity of other adults (Koford 1958).

Immigration and emigration factors appear 

to be related to prairie dog population 

fluctuations (Tileston and Lechleitner 1966). 

There have been few studies on dispersal 

and reported distances are usually maximum 

observations. Garrett and Franklin (1988) 

report an average black-tailed prairie dog 

dispersal distance of 2.4 km, but Koford 

(1958) observed black-tailed prairie dogs 

emigrating nearly 6.5 km. Clark (1973) 

recorded white-tailed prairie dogs moving 

2.7 km during emigration between March 

and April and between July and August. 

Hillman et al. (1979) reported that the 

mean distance between black-tailed prairie 

dog towns and their nearest neighbors 

was 2.4 km. As prairie dogs emigrate and 

establish new towns, they select habitat 

based on criteria that have not been fully 

defined. If habitat selection criteria were 

better defined, then efforts to conserve 

potential habitat might be more successful.

Habitat selection can vary greatly and might 

be explained by many factors, including 

environmental conditions, vegetation barriers, 

and prairie dog control efforts (Cambell III 

and Clark 1981; Coppock and Detling 1986; 

Weltzin et al. 1997). Data indicating prairie 

dog selection of certain soil and precipitation 

characteristics appears to be limited and 

information for the Bighorn Basin is 

needed. Other studies have observed new 

burrows in soils that are dense clay-clayeysaline uplands correlated with the cessation 

of annual plant growth or existing growth 

dehydration (Clark 1973; Parker et al. 1975).

The vegetative community and structure 

can also influence the distribution of prairie 

dogs (Osborn and Allan 1949; Weltzin et al. 

1997). Prairie dogs feed mainly on annual 

forbs and other plants typical of early 

successional stages, and their herbivory 

can increase shoot nitrogen concentrations 

(Bond 1945; Holland and Detling 1990). 

Their food habits vary with the time of year 

and the availability of vegetation (Tileston 

and Lechleitner 1966). Generally, blacktailed prairie dogs may eat mostly forbs and 

grasses, while the white-tailed prairie dogs 

may eat more grasses and sedges (Tileston 

and Lechleitner 1966). Prairie dog movement can be restricted in some cases by tall 

vegetation such as greasewood; however, 

white-tailed prairie dogs are more tolerant 

of taller vegetation than black-tailed (Clark 

1973; Hoogland 1996; Weltzin et al. 1997). 

Absence of prairie dogs can increase woody 

vegetation seed dispersal and alter species 

composition (Osborn and Allan 1949; 

Weltzin et al. 1997). Rosenstock and Van 

Riper III (2001) report active prairie dog 

towns were present in grasslands uninvaded 

by shrubs and absent in successional 

woodlands, indicating aversion to woody 

vegetation encroachment.

Prairie dog distribution and activity are 

dependent on many known and unknown 

biotic and abiotic attributes. The study area 

was defined and inventory methods were 

used in 2001–2005 that were similar to 

methods for the 1980–1989 BLM survey. 

This report describes trends in course scale 

population indices spanning 12 to 26 years, 

and also analyzes soil type and precipitation 

zone selection for the prairie dog population.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

3

THE PRAIRIE DOG TOWNS within the 

study area (Total Population) were divided 

into two metapopulations: the Basin 

Metapopulation and the Meeteetse 

Metapopulation. The Basin Metapopulation 

area was south of the Montana and 

Wyoming State line, east of the Shoshone 

National Forest, west of the Bighorn 

National Forest, and north of Shell Creek 

and the Greybull River (Figure 1). The 

Meeteetse Metapopulation area was 

defined as towns west of Highway 120, 

north of the Greybull River and the Wood 

River, east of the Shoshone National Forest 

and south of Latitude 44° 19’ 40.00” North 

(Figure 1). An observed white-tailed prairie 

dog dispersal distance of 2.7 km (Clark 1973) 

was used to separate the Meeteetse 

Metapopulation from the Basin 

Metapopulation.

STUDY AREA STUDY AREA

White-tailed prairie dog
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

Figure 1. The study area includes the Total Population, which is equal to the Basin Metapopulation plus the Meeteetse Metapopulation survey areas.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

5

THE 1980–1989 SURVEY was identified 

as including two major survey efforts, 

the 1980–1989 BLM survey data and the 

Meeteetse Metapopulation survey (mapped 

by the FWS on 1:100:000 mylar in 1988), 

which together surveyed the study area 

with similar land access issues and did 

not include lands up the South Fork of the 

Shoshone River. Active prairie dog town 

locations from these maps were screen 

digitized and areas were calculated in GIS 

Arc Map 9.1 ESRI 2005. Active 1980–1989 

towns were assigned a new consecutive 

town “ID” number by sorting through 

unknown and inactive towns. These 

numbers were linked to the original survey 

numbers, referenced in GIS and on data 

sheets in a new Prairie Dog Survey Binder 

and added to a database in Microsoft 

Office Access 2003. These data were then 

compared to a new survey conducted by the 

BLM between 2001 and 2005.

Because it took multiple years to accumulate 

data, the surveys are not instantaneous 

descriptions. Results represented active 

prairie dog towns found during the surveys, 

but surveys may not have been complete. 

The FWS survey did not include the same 

methods used by the 1980–1989 BLM 

survey; therefore, town burrow activity 

levels in the Meeteetse Metapopulation 

were not analyzed or compared to the 

2001–2005 survey.

In an effort to replicate the former 

survey with similar methods, surveys were 

conducted from April through November. 

Surveys were conducted on BLM, Bureau 

of Reclamation (BOR), state, and private 

lands when access was granted in potential 

prairie dog habitat, which included 

vegetation communities that were not 

in river bottoms, irrigated fields, forests, 

alpine, and subalpine areas. All previously 

identified prairie dog towns were resurveyed. 

Aerial photography and former survey 

results assisted in locating prairie dog 

towns. In addition to ground observations, 

local residents were interviewed to increase 

detection rates. Ridges and roads were 

driven or walked to maximize the visual 

area surveyed. The most effective method 

for finding prairie dog towns appeared 

to be binocular searches from elevated 

locations (Coy and Roberts 1985). Aircraft 

surveys are effective but were precluded 

due to expense. Once a town was identified, 

its location was recorded as a polygon 

using a global positioning system 

(Trimble GeoExplorer 3).

Town boundaries were defined by walking 

the perimeter to the closest burrow or 

mound, recording data with a GPS unit. A 

town was considered new if its boundaries 

did not overlap a town identified during a 

METHODS METHODS

Prairie rattlesnake occupying a black-tailed prairie dog burrow.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

6

previous survey. Once every data collection 

season on relatively high, medium, and 

low density towns, a measuring wheel was 

used to conduct 1 meter wide transects 

oriented north and south distributed every 

30 m parallel to each other continuing to 

the edges of the town to calibrate observer 

estimates for burrow density (Biggins et 

al. 1993). For the study, active and inactive 

burrow densities were estimated and placed 

within categories of ≤10 or >10 burrows 

per 0.405 ha (1.0 ac). A Prairie Dog Town 

Inventory Form was filled out according 

to the Cody Resource Area Prairie Dog 

Ecosystem Habitat Management Plan 

(Coy and Roberts 1985); the town was then 

assigned an identification number and 

incorporated into the database.

Data were analyzed for active towns within 

the Total Population and analyzed separately 

as the Meeteetse Metapopulation and 

the Basin Metapopulation (Figure 1). Any 

number of active burrows mapped at the 

finest resolution was considered a “town.” 

A town was considered active if there were 

any recent prairie dog signs of activity 

such as: observed individuals, fresh scat, 

vocalizations, and fresh excavation (Biggins 

et al. 2006). Data for inactive towns were 

not collected or analyzed during this survey; 

however, inactive burrows within an active 

town were used to estimate the inactive 

burrow density. Inactive prairie dog towns 

are present throughout the landscape, and 

they may be recently inactive or may have 

been inactive for many years (Biggins et al. 

2006). Although inactive towns are very 

important to wildlife still occupying the 

area, it could not be determined how long 

a town had been inactive and so they 

were not collected since they would not 

indicate current prairie dog distribution or 

abundance.

Nearest town distances for all 2001–2005 

white-tailed prairie dog towns were 

measured using GIS ArcMap 9.1 ESRI 2005. 

Summary statistics were produced for 

measured distances from the outer edge 

boundaries of all white-tailed prairie dog 

towns to the next nearest town edge. 

Nearest distances were averaged to 

produce the mean distance to the nearest 

neighboring town.

Soil type and precipitation zone selection 

analysis were based on “Actual” and 

“Potential” use. The 2001–2005 active 

white-tailed prairie dog town areas were 

analyzed as Actual use areas. A 2.7 km 

buffer area around the Actual use areas 

were analyzed as Potential use areas for 

white-tailed prairie dogs and it was assumed 

this buffer incorporated potential habitat 

for selection based on dispersal observations 

(Clark 1973).

In GIS, Actual and Potential areas were 

overlaid on top of Park and Bighorn 

County’s soil surveys, which also included 

precipitation zone data that are digitized, 

initial field work performed by the BLM 

and the Soil Conservation Service, now the 

Natural Resources Conservation Service 

(NRCS), in the late 1970s and early 1980s. 

Total soil type areas and precipitation areas 

were calculated within the Actual and 

Potential areas.

The resulting soil type areas were divided 

into range site areas using the percent 

occurrence listed in the map unit. For 

instance, a town was found to have 

247.1 ha (610.0 ac) of soil type 474AD, 

which generally has range sites with 

40 percent shale, 50 percent saline upland, 

and 10 percent that is “not specified.” 

Therefore, the area was proportioned by 

percentage and it was assumed that 98.7 ha 

(244.0 ac) were used in shale sites, 123.4 ha 

(305.0 ac) were in saline uplands, and 24.7 ha 

(61.0 ac) were not named. Unspecified soil 

types were excluded from the analysis.

Area and percent occurrence of each 

precipitation zone were also calculated. 

Using the five most frequent range sites and 

the three precipitation zones, which overlap 

[12.7–22.9 cm (5–9 in), 20.3–25.4 cm 

(8–10 in), and 25.4–35.6 cm (10–14 in)]; 

a two-dimensional chi square test was 

performed between the Actual and the 

Potential available area (2.7 km buffer). This 

method assumes all range sites within a soil 

type are used equivalently as often as they 

occur and that soil surveys for private lands 

were not entirely complete.

Descriptive data analyses were primarily 

conducted in Microsoft Office Excel 2003. A 

two-dimensional chi-square test was used 

to test for significant associations between 

the two surveys and the two categories of 

burrow density, the two categories of 

activity level, precipitation zone area, 

and range site area within 2.7 km of the 

white-tailed prairie dog town edge and the 

area within towns. The Frequency unit was 

per hectare for the precipitation and range 

site statistical analysis. Null hypotheses 

were rejected at α = 0.05. Statistical tests 

were performed in Microsoft Excel 2003 

and in VassarStats: Web Site for Statistical 

Computation (Lowry 2007).
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

7

Population

Demographics

The 1980–1989 survey results 

for the Total Population:

There were 105 active white-tailed towns 

occupying 7,031.8 ha (17,374.9 ac) 

with a minimum area of 0.008 ha 

(0.02 ac) and a maximum of 1,607.4 ha 

(3,972.0 ac). The mean area was 66.4 ha 

(164.0 ac) with a 178.3 ha (440.5 ac) 

standard deviation. Two towns were 

black-tailed prairie dog towns totaling 

71.4 ha (176.4 ac).

The 1980–1989 survey results for the 

Meeteetse Metapopulation:

There were 49 active white-tailed towns 

occupying 4,736.7 ha (11,704.6 ac) with 

a minimum area of 0.04 ha (0.1 ac) and a 

maximum of 1,607.4 ha (3,972.0 ac). The 

mean area was 96.7 ha (239.0 ac) with a 

242.6 ha (599.5 ac) standard deviation.

The 1980–1989 survey results for the 

Basin Metapopulation:

There were 56 active white-tailed towns 

occupying 2,294.7 ha (5,670.3 ac) with a 

minimum area of 0.008 ha (0.02 ac) and 

a maximum of 631.4 ha (1,560.1 ac). The 

mean area was 40.8 ha (100.8 ac) with 

an 89.9 ha (222.2 ac) standard deviation. 

Two towns were black-tailed prairie dog 

towns totaling 71.3 ha (176.4 ac).

The 2001–2005 survey results for the 

Total Population:

There were 66 active white-tailed towns 

occupying 2,011.5 ha (4,970.6 ac) with a 

minimum area of 0.08 ha (0.2 ac) and a 

maximum of 572.1 ha (1,413.8 ac). The 

mean area was 29.8 ha (73.7 ac) with a 

75.6 ha (186.7 ac) standard deviation. 

Four towns were black-tailed prairie dog 

towns totaling 77.4 ha (191.3 ac), an 

increase of 8 percent.

The 2001–2005 survey results for the 

Meeteetse Metapopulation:

There were 5 active white-tailed towns 

occupying 615.2 ha (1,520.1 ac) with a 

minimum area of 1.0 ha (2.5 ac) and a 

maximum of 572.1 ha (1,413.8 ac). The 

mean area was 123.0 ha (304.0 ac) with 

a 251.3 ha (621.0 ac) standard deviation. 

RESULTS RESULTS

White-tailed prairie dog
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

8

The 2001–2005 survey results 

for the Basin Metapopulation:

There were 61 active white-tailed towns 

occupying 1,396.4 ha (3,450.5 ac) with 

a minimum area of 0.08 ha (0.2 ac) and 

a maximum of 271.7 ha (671.5 ac). The 

mean area was 22.7 ha (56.0 ac) with 

a 38.4 ha (94.9 ac) standard deviation. 

Four towns were black-tailed prairie dog 

towns totaling 77.4 ha (191.3 ac).

The mean nearest neighbor distance 

between white-tailed towns for the 

2001–2005 survey was 2,013 m, with a 

minimum of 60 m, maximum of 9,057 m, 

and a standard error of 279.7. The mean 

nearest distance between towns for the 

1980–1989 survey was 1,963 m, with a 

minimum of 48 m, maximum of 13,243 m, 

and a standard error of 235.7. The whitetailed prairie dog population also comprised 

96 percent of the occupied prairie dog 

area and 4 percent of the area was 

occupied by black-tailed prairie dogs 

for the 2001–2005 survey.

The percent decrease in occupied area 

occurring between the 1980–1989 surveys 

and the 2001–2005 survey was 71 percent 

for the Total Population, 87 percent for the 

Meeteetse Metapopulation, and 39 percent 

for the Basin Metapopulation (Table 1 

and Figure 2). The 2001–2005 Basin 

Metapopulation and Total Population had 

a smaller active mean town area than the 

1980–1989 Total Population (Table 1). The 

2001–2005 Meeteetse Metapopulation 

mean town area was larger than the 

1980–1989 Meeteetse Metapopulation.

Figure 2. Comparison of hectares occupied by white-tailed prairie dogs for the Total Population, 

Meeteetse Metapopulation, and Basin Metapopulation during the two surveys beginning in 

1980 and 2001.

Table 1. Attributes of white-tailed prairie dog towns surveyed during 1980–1989 and 2001–2005. 

1980–1989 Survey 2001–2005 Survey Percent Difference

Total Population

Town Abundance 105 66 37 percent decrease

Occupied Area 7,031.8 ha (17,374.9 ac) 2,011.5 ha (4,970.6 ac) 71 percent decrease

Mean Town Area 66.4 ha (164.0 ac) 29.8 ha (73.7 ac) 55 percent decrease

Meeteetse Metapopulation

Town Abundance 49 5 90 percent decrease

Occupied Area 4,736.7 ha (11,704.6 ac) 615.2 ha (1,520.1 ac) 87 percent decrease

Mean Town Area 96.7 ha (239.0 ac) 123.0 ha ( 304.0 ac) 79 percent increase

Basin Metapopulation

Town Abundance 56 61 8 percent increase

Occupied Area 2,294.7 ha (5,670.3 ac) 1,396.4 ha (3,450.5 ac) 39 percent decrease

Mean Town Area 40.8 ha (100.8 ac) 22.7 ha (56.0 ac) 44 percent decrease
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

9

Burrow density and activity within the 

Basin Metapopulation appears to have 

changed over time (Figures 3 and 4; Table 2). 

There was a trend towards more inactive 

burrows in both the ≤10 and >10 burrows 

Figure 3. Basin Metapopulation frequency of active and inactive 

burrow density estimates within active white-tailed prairie dog 

towns categorized as ≤10 or >10 burrows per 0.405 hectare 

(1.0 ac) for the 1980–1989 survey.

Figure 4. Basin Metapopulation frequency of active and inactive 

burrow density estimates within active white-tailed prairie dog 

towns categorized as ≤10 or >10 burrows per 0.405 hectare 

(1.0 ac) for the 2001–2005 survey.

Table 2. Basin Metapopulation percent change 

during the time between the two surveys of 

active and inactive burrows within active white 

tailed prairie dog towns categorized as ≤ 10 or 

>10 burrows per 0.405 ha (1.0 ac) categories.

Active Inactive

≤10 -32 percent +100 percent

>10 -33 percent +71 percent

per 0.405 ha (1.0 ac) categories for the 

2001–2005 survey and were significantly 

associated with the time since the 

1980-1989 survey (X2 = 12.76, P < 0.006, 

Cramer’s V = 0.24) (Figures 3 and 4; Table 2).

Habitat Associations

A two-dimensional chi-square test of the 

five most frequent range sites showed a 

significant association between the actual 

range site areas selected by white-tailed 

prairie dogs when compared to potential 

dispersal site areas available within the 

potential dispersal area (2.7 km) (X2 = 77.67, 

P < 0.0001 and Cramer’s V = 0.017)(Figure 5). 

Of the soil types present in active whitetailed prairie dog towns, saline uplands 

and loamy sites were often a major soil 

component and were most frequently used. 

They made up 32.6 percent and 22.8 

percent of the total active white-tailed 

prairie dog town area respectively (Figure 5).

Saline upland and loamy sites appear to be 

associated with white-tailed prairie dog 

distribution, as soils in the potential dispersal 

area were relatively less frequent (Figure 5). 

The two largest differences between the 

actual areas selected and the potential areas 

available were in clayey and shale range 

sites. This illustrates a higher frequency of 

use of these sites than the amount theoretically available to them. Saline uplands and 

loamy sites were most common; however, 

white-tailed prairie dogs appear to have 

a high selective propensity for infrequent 

clayey and shale range sites. There appears to 

be some selection for loamy sites; however, 

for shallow loamy, the actual and potential 

percent selection was identical (Figure 5).
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

10

Figure 5. Area and percent of range site area actually occupied by 2001–2005 

active white-tailed prairie dog towns compared with potentially occupied areas 

within 2.7 km.

There also appears to be a significant 

association between precipitation zones 

and white-tailed prairie dog distribution 

(X2 = 928.89, P < 0.0001 and 

Cramer’s V = 0.047). The only zone where 

the active town occurrence was greater 

than what was available was in 

the 20.3–25.4 cm (8–10 in), precipitation 

zone (Figure 6). Thus, range sites and 

precipitation zones account for part of the 

observed variability for site selection. Low 

Cramer’s V values indicate other variables 

also influence white-tailed prairie dog town 

site selection.

Figure 6. Area of land in precipitation zones 12.7–22.9 cm (5–9 in), 20.3–25.4 cm (8–10 in), and 

25.4–35.6 cm (10–14 in), occupied by 2001–2005 active white-tailed prairie dog towns and 

potentially occupied areas within 2.7 km. 

White-tailed prairie dog
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

11

THERE APPEARS TO HAVE BEEN MANY 

CHANGES IN PRAIRIE DOG ACTIVITY and 

distribution during the relatively short time 

period (12–26 year range) between the 

two surveys. Within the study area, blacktailed prairie dog towns appear to have 

increased slightly, while the Total Population 

of white-tailed prairie dog towns have 

decreased as a whole, showing large 

decreases in four categories: town 

abundance, occupied area, mean town 

area, and active burrow density (Tables 1 

and 2; Figures 2, 3, and 4). In other words, 

active white-tailed prairie dog towns were 

fewer, smaller, and more inactive.

Although the black-tailed prairie dog 

population increased slightly, it may be 

vulnerable since they are so rare and 

isolated from the rest of the species’ range. 

White-tailed prairie dogs are by far the 

most common prairie dog species in the 

study area, which is probably due to their 

adaptations for sagebrush vegetation 

communities (Hoogland 1996).

The two white-tailed prairie dog metapopulations (Meeteetse Metapopulation 

and Basin Metapopulation) within the Total 

Population appear to have changed in 

different ways. The Meeteetse Metapopulation, where black-footed ferrets 

were found, showed the largest decline. 

There are fewer, but larger towns, covering 

much less area (a large decrease in occupied 

area and town abundance, and an increase 

in the mean town area - Table 1). This 

suggests that surviving white-tailed prairie 

dogs may have contracted to core areas, 

possibly resulting in part from epizootic and 

enzootic plague, along with many other 

cumulative factors (Menkens and Anderson 

1991; Cully and Williams 2001). Currently 

this metapopulation has become much less 

suitable for black-footed ferrets, although 

it may rebound to a suitable size where 

ferrets could one day be returned to where 

the recovery source population was found.

The Basin Metapopulation appears to have 

become more fragmented. There are more, 

but smaller towns, which cover less area 

(occupied area and mean town area both 

decreased, while the town abundance 

increased - Table 1). Within the Basin 

Metapopulation, white-tailed prairie dogs 

may be more easily fragmented than the 

Meeteetse Metapopulation due to highways, 

urban areas, and other development 

interspersed throughout the population.

Within the Basin Metapopulation, there 

was also a significant association between 

categorical estimates of active and inactive 

burrow densities between the 1980–1989 

BLM surveys and the 2001–2005 survey. 

DISCUSSION DISCUSSION

Cottontail rabbit occupying an inactive white-tailed prairie dog burrow.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

12

Comparisons of burrow density could not 

be made in the Meeteetse Metapopulation 

because data were not collected with the 

same methods. These results suggest that 

during the time between the two surveys, 

there must have been certain factors causing 

a shift from relatively higher to lower density 

estimates of active burrows and an increase 

in inactive burrow density estimates 

(Figures 3 and 4). Causal factors of this 

shift and the other observed demographic 

declines were not analyzed and may be any 

of the identified causes for declines (Cully 

and Williams 200; Pauli et al. 2006; Seglund 

et al. 2004); however, enzootic plague may 

be a plausible explanation since plague 

has been detected every year sampled in 

the Meeteetse Metapopulation by the U.S. 

Geological Survey from 1988–2008 (Dean 

Biggins per. comm.).

Trends in these colony attributes are the 

best available indicators of prairie dog 

population trends in the northern Bighorn 

Basin. Thus, considering the overall decrease 

in occupied area and a proportional shift 

from more active burrows to more inactive 

burrows, the white-tailed prairie dog 

population within this study area 

appears to have exhibited negative trends 

in population distribution and abundance 

(Lomolino and Smith 2001). Efforts to 

reverse these trends should be implemented 

before further decreases occur, which could 

further reduce the overall white-tailed 

prairie dog range and distribution. Another 

short-term decline could lead to additional 

justification for listing under the Endangered 

Species Act of 1973.

These results should be interpreted 

cautiously due to potential biases and 

limitations of the study design. It is quite 

difficult to accurately and precisely replicate 

historical surveys where the goal is to 

inventory prairie dog distribution and 

activity (Seglund et al. 2004). This study 

assumes that all towns in the study area 

were included in this analysis, or at least 

that the levels of detection were the same 

for both surveys. The results of both surveys 

are likely underestimated, biased towards 

prairie dog towns on Federal and Stateadministered lands, and larger towns. 

Because access to private land was not 

possible in all cases, there may be prairie 

dog towns that were not detected during 

the surveys.

In order to compare 12–26 year old 

survey data to the present survey data, 

the 1980–1989 surveys methods were 

replicated and limitations for both surveys 

were assumed to be similar. Access to land, 

however, may have changed between the 

two surveys because of changing land 

ownership patterns. GPS units were used 

to map towns, which may have been more 

accurate than hand-drawn maps from the 

previous surveys.

Although visual counts might provide 

more accurate indices to actual populations 

(Severson and Plumb 1998), the methods 

used for this study were designed to 

replicate the former study and time and 

resources were limited. The results should 

be appropriate for trend estimates on 

activity, occupied area, and distribution for 

the study area (Biggins et al. 1993, 2006; 

Lomolino and Smith 2001). Attempting to 

replicate a historical survey by using similar 

methods has long-term trend analysis 

value and may be impractical to use more 

rigorous methods (e.g., visual counts, 

mark-recapture, and mark-resight on 

large-scale studies) (Biggins et al. 2006).

Although there are errors with estimating 

burrow density, the error was reduced by 

using categories large enough where 

estimates fall within a low rating of ≤10 or 

a high rating of >10 inactive or active 

burrows per 0.405 hectare (1.0 ac). Calibrating 

observers to estimate high and low burrow 

densities seems to be a reasonable approach 

for estimating activity level as categorical 

frequency data. If there were more resources 

and time, burrow density transects could 

have been deployed on every prairie dog 

town providing for more accurate and 

precise burrow density and activity data.

The habitat selection phase of this study 

was aimed at describing the spatial 

distribution of towns relative to soils, 

precipitation, and proximity to the nearest 

town. An area with a perimeter boundary 

of 2.7 km from all surveyed towns was used 

for a potential dispersal distance and seems 

appropriate for comparisons of range site 

and precipitation zone habitat selection 

by white-tailed prairie dogs (Clark 1973). 

The mean minimum distance that was 

measured between towns (2 km) was less, 

though similar to observations of Clark 

(1973) (2.7 km) and Garrett and Franklin 

(1988) (2.4 km). The mean minimum 

distance only describes white-tailed prairie 

dog town spatial distribution, which may 

be related to many factors including habitat 

quality, individual dispersal behavior, and 

habitat fragmentation (Clark 1973; Cambell 

III and Clark 1981; Coppock and Detling 

1986; Weltzin et al. 1997).

Considering the observed declines in this 

population, minimizing disturbances and 

managing for possible town expansion 

within a 2 km buffer zone from prairie dog 

towns may help maintain current dispersal 

and prairie dog town connectivity. Buffers 

around prairie dog towns may need to be 

adjusted depending upon the disturbance 

type and local habitat variation. Future 
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

13

surveys could measure the mean minimum 

distance again and use it as an indicator for 

how distribution may be trending.

White-tailed prairie dog town sites were 

significantly associated with certain range 

sites and precipitation zones. This study’s 

results showed an association with multiple 

range sites including: saline upland, loamy, 

shale, and clayey sites. Parker et al. (1975) 

observed similar sites including dense 

clay-clayey-saline upland selection. Other 

variables also likely have an effect on these 

town locations as the Cramer’s V is relatively 

low (Cramer’s V = 0.017).

Other habitat selection variables that may 

be contributing to declines and distribution 

may include: range site condition, vegetation 

and watershed health, soil permeability, 

and the presence or absence of other 

wildlife species (Garrett et al. 1982; Ceballos 

and Reading 1994). Sylvatic plague may be 

the number one factor, which frequently 

kills >99 percent of prairie dogs in infected 

colonies (Cully and Williams 2001). Mortality 

through shooting or poisoning and 

habitat loss may also be contributing to the 

observed declines. All these factors should 

be studied to explain the cumulative effects 

contributing to the observed population 

decline in the northern Bighorn Basin and 

throughout their range.

This study shows that saline upland, loamy, 

sandy, shallow loamy, and shale range 

sites were significantly associated with 

white-tailed prairie dog town site selection 

when compared to available habitat within 

2.7 km (Figure 5). White-tailed prairie dogs 

occupied the most area in the 12.7–22.9 cm 

(5–9 in) precipitation zone and in the saline 

uplands range site; however, loamy sites and 

the 20.3–25.4 cm (8–10 in) precipitation 

zone have the highest proportion of actual 

active area than what is proportionally 

potentially available within the 2.7 km 

buffer (Figures 5 and 6). This suggests that 

white-tailed prairie dogs may be more 

successful within this range site and 

precipitation zone. Also, available range 

sites less suitable for white-tailed prairie 

dog towns, such as rock outcrops and 

impervious clays, were nearly absent from 

the actual soils used and could be limiting 

dispersal and site suitability (Garrett et al. 

1982). The range site and precipitation 

zone analyses should be considered a 

broad scale assessment from soil survey 

information, using overlapping precipitation 

zone categories, and may not reflect unique 

diversity in local conditions. However, these 

results may help to refine existing habitat 

suitability models.

Soil types may also be chosen by whitetailed prairie dogs for their structural 

components (Flath and Paulick 1979). 

Many soils could be difficult to tunnel in 

and inhibit dispersal. A shallow soil may 

not allow enough depth to bedrock, soils 

of impervious clays would be very difficult 

to dig in, being directly in a wetland or 

floodplain could mean many months of 

inundation, and sandy soils lacking any other 

matrix could be prone to collapse (Pauli et 

al. 2006). Soils with high percentages of 

clay would lack permeability, which could 

increase flooding and standing water in 

burrows. This study’s results suggest and 

it would seem reasonable that a loamy 

soil would be most suitable, as tunnels 

may require a proportion of clay to hold 

the walls, yet the addition of sand and silt 

would allow for easier burrowing.

Since prairie dogs can have higher survivorship in areas with more vigorous forbs, 

grasses, and sedges, historically they may 

have built their towns on more productive 

loamy sites, possibly those which are 

now primarily agricultural tracts (Garrett 

et al. 1982). It is hard to say how much 

present land conditions and uses prohibit 

prairie dogs from expanding their range. 

However, practices such as grazing and 

agriculture regularly occur on the most 

productive soils and have displaced prairie 

dog towns (Koford 1958). White-tailed 

prairie dogs currently occupy soils, whose 

alkalinity, texture, lack of depth, and lower 

precipitation are not generally converted 

to agriculture. These less productive sites 

may not adequately provide all life history 

requirements and perhaps may be all the 

suitable land that is left to occupy in a fairly 

undeveloped or seasonally undisturbed 

state (Garrett et al. 1982). This survey shows 

current soil selection and may not represent 

historical, undisturbed habitat selection. 

Studies of town locations before settlement 

would illustrate which range sites were 

used in a less disturbed ecosystem; 

however, historical evidence may be 

uncommon and is likely eroding fast.

Soils and precipitation appear to have an 

effect on habitat selection by prairie dogs. 

Displacement from these preferred areas 

may contribute to the observed decline 

within the study area. There are surely 

other habitat and anthropogenic variables 

influencing the prairie dog decline. Plague 

epizootics caused by flea-transmitted 

bacteria and recreational shooting can 

substantially affect prairie dog town 

persistence (Lechleitner et al. 1968; Pauli 

et al. 2006; Pauli and Buskirk 2007) and it 

is likely that plague has had overwhelming 

influence on prairie dog population dynamics 

(Dean Biggins per. comm.). Investigations 

on how natural habitat selection is 

influencedby anthropogenic factors should 

be studied further and may reveal additive 

effects on normal population fluctuations.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

14

Reversing the observed negative population 

trend is important for conserving a species 

that provides a prey base for a complex 

ecosystem and habitat for species, which 

are BLM sensitive species or listed under the 

Endangered Species Act of 1973, such as: 

black-footed ferret, burrowing owl (Athene 

cunicularia), mountain plover (Charadrius 

montanus), and ferruginous hawk (Buteo 

regalis) (Bangert and Slobodchikoff 2000). 

This is a short and incomplete species list 

for an ecosystem dependent upon the 

presence and distribution of prairie dogs 

(Campbell and Clark 1981; Ceballos and 

Reading 1994; Miller et al. 1994; 

Desmond and Savidge 1996; Bangert 

and Slobodchikoff 2000).
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

15

Bond, R.M. 1945. Range rodents and plant 

succession. Trans North American Wildlife 

Conference 10:229-234.

Campbell III, T.M. and T.W. Clark. 1981. 

Colony characteristics and vertebrate 

associates of white-tailed and blacktailed prairie dogs in Wyoming. American 

Midland Naturalist 105(2):269-276.

Cambell III, T.M., T.W. Clark, L. Richardson, 

S.C. Forrest, and B.R. Houston. 1987. 

Food habits of Wyoming black-footed 

ferrets. American Midland Naturalist 

117(1):208-210.

Clark, T.W. 1973. A field study of the ecology 

and ethology of the white-tailed prairie 

dog (Cynomys leucurus): with a model of 

Cynomys evolution. Ph.D. dissertation, 

University of Wisconsin, Madison.

Clark, T.W. and M.R. Stromberg. 1987. 

Mammals in Wyoming. University Press 

of Kansas. Lawrence, Kansas, USA.

Coppock, D.L. and J.K. Detling. 1986. 

Alteration of bison and black-tailed 

prairie dog grazing interaction by 

prescribed burning. Journal of Wildlife 

Management 50(3):452-455.

Coy, S.G. and D.A. Roberts. 1985. Prairie dog 

ecosystem habitat management plan. 

USDI Bureau of Land Management. No. 

WY-010-WHA-T14.

Cully, J.F. and E.S. Williams. 2001. Interspecific comparisons of Sylvatic plague 

in prairie dogs. Journal of Mammalogy 

82(4):894-905. 

Desmond, M.J. and J.A. Savidge. 1996. 

Factors influencing burrowing owl 

(Speotyto cunicularia) nest densities and 

numbers in western Nebraska. American 

Midland Naturalist 136(1):143-148.

Flath, D.L. and R.K. Paulick. 1979. Mound 

characteristics of white-tailed prairie dog 

maternity burrows. American Midland 

Naturalist 102(2):395-398.

Bangert, R.K. and C.N. Slobodchikoff. 2000. 

The Gunnison’s prairie dog structures a 

high desert grassland landscape as a 

keystone engineer. Journal of Arid 

Environments 46:357-369.

Biggins, D.E., B.J. Miller, L.R. Hanebury, 

B. Oakleaf, A.H. Farmer, R. Crete, and A. 

Dood. 1993. A technique for evaluating 

black-footed ferret habitat. Proceedings 

of the Symposium on the Management 

of Prairie Dog Complexes for the 

Reintroduction of the Black-footed 

Ferret. USDI Fish and Wildlife Service. 

Biological Report 13:73-88.

Biggins, D.E., J.G. Sidle, D.B. Seery, and A.E. 

Ernst. 2006. Estimating the abundance 

of prairie dogs. In: Hoogland, J.L., (ed.). 

Conservation of the black-tailed prairie 

dog: saving North America’s western 

grasslands. Washington, DC: Island Press. 

p. 94-107.

LITERATURE CITED LITERATURE CITED

Black-tailed prairie dog, Brian Holmes, Colorado Division of Wildlife
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

16

Lehmer, E.M. and D.E. Biggins. 2005. 

Variations in torpor patterns of freeranging black-tailed and Utah prairie 

dogs across gradients of elevation. 

Journal of Mammalogy 86:15-21.

Lomolino, M. and G.A. Smith. 2001. 

Dynamic biogeography of prairie dog 

(Cynomys ludovicianus) towns near 

the edge of their range. Journal of 

Mammalogy 82(4):937-945.

Lowery, R. 2007. Vassarstats: website for 

statistical computation. http://faculty.

vassar.edu/lowry/VassarStats.html

(accessed Feb. 25, 2007).

Menkens, G.E., Jr. 1987. Temporal and 

spatial variation in white-tailed prairie 

dog (Cynomys leucurus) populations 

and life histories in Wyoming. Ph.D. 

dissertation, University of Wyoming, 

Laramie, USA.

Menkens, G.E., Jr. and S.H. Anderson. 1991. 

Population dynamics of white-tailed 

prairie dogs during an epizootic of 

sylvatic plague. Journal of Mammalogy 

72(2):328-331.

Menkens, G.E., Jr. and S.H. Anderson. 1989. 

Temporal-spatial variation in whitetailed prairie dog demography and life 

histories in Wyoming. Canadian Journal 

of Zoology 67:343-349.

Miller, B., G. Ceballos, and R. Reading. 1994. 

The prairie dog and biotic diversity. 

Conservation Biology 8(3):677-681.

Osborn, B. and P. Allan. 1949. Vegetation of 

an abandoned prairie-dog town in tall 

grass prairie. Ecology 30(3):322-332.

Parker, J.L., G.L. Decker, L. Gray, and O. 

Muller. 1975. Soil survey of Carbon 

County area, Montana. U.S. Department 

of Agriculture, Soil Conservation Service. 

137 pp.

Pauli, J.N., S.W. Buskirk, E.S. Williams, and 

W.H. Edwards. 2006. A plague epizootic 

in the black-tailed prairie dog (Cynomys 

ludovicianus). Journal of Wildlife Diseases 

42(1):74-80.

Pauli, J.N. and S.W. Buskirk. 2007. 

Recreational shooting of prairie dogs: 

a portal for lead entering wildlife food 

chains. Journal of Wildlife Management 

71(1):103-108.

Rosenstock, S.S. and C. Van Riper III. 2001. 

Breeding bird responses to juniper 

woodland expansion. Journal of Range 

Management 54(3):226-232.

Seglund, A.E., A.E. Ernst, M. Grenier, B. 

Luce, A. Puchniak, and P. Schnurr. 2004. 

White-tailed prairie dog conservation 

assessment. Unpublished Report.

Severson, K.E. and G.E. Plumb. 1998. 

Comparison of methods to estimate 

population densities of black-tailed 

prairie dogs. Wildlife Society Bulletin 

26(4):859-866.

Tileston, J.V. and R.R. Lechleitner. 1966. 

Some comparisons of the black-tailed 

and white-tailed prairie dogs in northcentral Colorado. American Midland 

Naturalist 75(2):292-316.

Weltzin, J.F., S. Archer, and R.K. 

Heitschmidt. 1997. Small-mammal 

regulation of vegetation structure 

in a temperate savanna. Ecology 

78(3):751-763.

Garrett, M.G. and W.L. Franklin. 1988. 

Behavioral ecology of dispersal in the 

black-tailed prairie dog. Journal of 

Mammology 69(2):236-250.

Garrett, M.G., J.L. Hoogland, and W.L. 

Franklin. 1982. Demographic differences 

between an old and a new colony of 

black-tailed prairie dogs (Cynomys 

ludovicianus). American Midland 

Naturalist 108(1):51-59

Harlow, H.J. and Menkens G.E., Jr. 1986. A 

comparison of hibernation in the blacktailed prairie dog, white-tailed prairie 

dog, and Wyoming ground squirrel. 

Canadian Journal of Zoology/Revue 

Canadienne de Zoologie 64(3):793-796.

Hillman, C.N., R.L. Linder, and R.B. 

Dahlgren. 1979. Prairie dog distribution 

in areas inhabited by black-footed 

ferrets. American Midland Naturalist 

102(1):185-187.

Holland, E.A., and J.K. Detling. 1990. 

Plant response to herbivory and 

belowground nitrogen cycling. Ecology 

71(3):1040-1049.

Hoogland, J.L. 1996. Cynomys ludovicianus. 

Mammalian Species 535:1-10.

Hoogland, J.L. 2001. Black-tailed, Gunnison’s 

and Utah prairie dogs reproduce slowly. 

Journal of Mammalogy 82(4):917-927.

Knowles, C.J., C.J. Stoner, and S. P. Gieb. 

1982. Selective use of black-tailed prairie 

dog towns by mountain plovers. Condor 

84:71-74.

Koford, C.B. 1958. Prairie dogs, whitefaces, 

and blue grama. Wildlife Monographs 

3:1-78.

Lechleitner, R.R., L. Kartman, M.I. 

Goldenberg, and B.W. Hudson. 1968. An 

epizootic of plague in Gunnison’s prairie 

dogs (Cynomys gunnisoni) in southcentral Colorado. Ecology 49(4):734-743.
Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming

17

The mention of company names, trade names, or commercial products does not constitute endorsement or recommendation for use by the Federal Government.