Habitat Selection and Changes in the White-tailed and Black-tailed Population within the Northern Bighorn Basin, Wyoming BLM/WY/ ST-09/031+1110 www.blm.gov/wy/st/en/field_offices/Cody.html Available online at: Denver, Colorado 80225 P.O. Box 25047 Section Information andPublishing Services National Operations Center Bureau ofLand Management provided by: services Production Harrell, D., andL.Marks. andchangesinthewhite-tailed 2009.Habitat selection Citation: Suggested Harrell,Destin BLM Cody Field Office Cover photoprairie dog)andallotherphotos (white-tailed except asnoted by: Management, Cody Field Office, Wyoming. BLM/WY/ ST-09/031+1110. 16 pp. Wyoming, Technical Note 431.U.S. oftheInterior. Department Bureau ofLand and black-tailed Basin, Bighorn prairie dogpopulation withinthenorthern

Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming White-tailed prairie dog prairie White-tailed By: Destin Harrell Biologist Wildlife Management of Land Bureau Wyoming Office, Field Cody Lisa Marks Geologist Management of Land Bureau Wyoming Office, Field Cody 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 Black-tailed prairie dog, Randy Hayes, BLM Hayes, Randy dog, prairie Black-tailed

and advice

. Bill Wilson was always there for for there always was Wilson Bill .

include Brian Latta, Bill Palmeroy, Katie Katie Bill Palmeroy, include Brian Latta, Biggins, Dean Stafford. and Darci VinZant, and Martin BobDennis Oakleaf, Saville, much helpful offered Grenier was Richard Lowry with statistical questions. questions. Lowry with statistical Richard was enough kind members who were crew Field this publication collect for to much of the data review Stewart Mike processing. GPS and GIS data ideas, reviews, in offering generous was Rohling and Janine and support. Kathy Center Operations the National at Koselak effortsby editing, invaluable contributed original artwork and drawing for arranging, Bureau the to thanks finally, And this paper. Office Field Cody Management’s Land of for helping with this staff and management effort. collaborative WE WOULD LIKE TO THANK Joe Capodice, THANK Joe Capodice, TO LIKE WOULD WE helped the BLM. He biologist for retired Kiracofe Steve survey. dog prairie our inspire the soil survey a big help for was analysis as

ACKNOWLEDGMENTS ACKNOWLEDGMENTS 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 Burrowing owls Burrowing 1 3 5 7 7 9 vii 11 15 ......

......

Population Demographics Population Associations Habitat ABSTRACT INTRODUCTION LITERATURE CITED LITERATURE STUDY AREA STUDY METHODS RESULTS DISCUSSION

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

approximately 2.0 km. Towns appear to be to appear Towns 2.0 km. approximately with saline upland, associated significantly and shale loamy, shallow sandy, loamy, prairie White-tailed 0.0001). < (P sites range dogs were also significantly associated associated also significantly dogs were and (P < 0.0001) zones with precipitation the 20.3–25.4 cm (8–10 in) prefer may though they even zone precipitation in the 12.7–22.9 cm common more were Other variables zone. (5–9 in) precipitation the to contribute also likely not measured observed distribution of white-tailed prairie selection. site dog town decrease in the mean active town area. area. in the mean active town decrease the Meeteetse for Declines area occupied in greater were (87 percent) Metapopulation population remaining the for declines than There - 39 percent). (Basin Metapopulation also a very of black- was small population that ludovicianus) dogs (C. tailed prairie 77.4 ha (191.3 ac). to 8 percent increased declined Density of active burrows for the (P < 0.006) significantly Basin mean nearest The Metapopulation. white-tailed separating neighbor distances both surveys for were dog towns prairie

WITHIN THE NORTHERN BIGHORN BASIN, THE NORTHERN WITHIN survey dog (Cynomys) prairie Wyoming, were 1989 and 1980 between collected data compared with data from a 2001–2005 from with data compared distribution, these data, From survey. selection and habitat activitybased level, categories and precipitation site on range the white-tailed Overall, analyzed. were declined population leucurus) dog (C. prairie Occupied active in distribution and activity. 7,031.8 ha from 71 percent decreased area ac); 2,011.5 ha (4,970.6 (17,374.9 ac) to in decrease also a 37 percent was there and a 55 percent abundance active town ABSTRACT

ABSTRACT

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 Mountain plover, endemic to North America, find prairie dog towns to be good nesting habitat. nesting good be to towns dog prairie find America, North to endemic plover, Mountain

with and adapted to prairie dog towns dog towns prairie to with and adapted and Clark 1979; Campbell (Hillman et al. During 1987). the et al. 1981; Cambell 26, 1981, a black- of September evening dog a rancher’s by killed was ferret footed (30 miles) south 48 km approximately of west kilometers and a few of Cody the consequential Wyoming; Meeteetse, extinct thought of the once capture live 1981 October in occurred ferret black-footed 1991). In 1985, and Anderson (Menkens pestis) was plague (Yersinia sylvatic metapopulation dog prairie the in discovered of the population where near Meeteetse discovered was ferrets black-footed 1991). and Anderson (Menkens James prompt helped to discoveries These of Land of the Bureau Coy and Steven Bredy (Mustela nigripes) (Mustela ferret black-footed The the majority dogs for upon prairie of preys associated species many of one is and diet its

and Clark 1981; Knowles et al. 1982; Miller Miller 1982; et al. 1981; Knowles and Clark 1997). et al. Weltzin 1994; et al. prairie dog range is unknown. Speculation Speculation is unknown. dog range prairie isolated an or captivity from release includes range. of their former population dogs and black-tailed White-tailed prairie and histories life different dogs have prairie in different the ecosystem influence may clip dogs may White-tailed prairie ways. prairie less than black-tailed vegetation white- Duringwinter, 1973). the (Clark dogs an at torpor enter dogs may tailed prairie longer of 7° C for air temperature ambient prairie periods of time than black-tailed hibernators facultative which are dogs, 1986; and Menkens (Harlow during winter and Biggins 2005). Both species Lehmer unique providing their environment, alter and can be a species, associated for habitat (Campbell predators for base prey substantial

, are often often , are spp.) (Cynomys

PRAIRIE DOGS described as a keystone species and described as a keystone 1994; engineers (Millerecological et al. Bangert 2000), have and Slobodchikoff throughout declined as much as 98 percent settlement North European America since of Viability and Reading 1994). (Ceballos an important species has become several or listed species are All issue. biodiversity listing under the been petitioned for have Species Act of 1973. Endangered in the issue is relevant biodiversity This white- where Wyoming, Bighorn Basin, near the are leucurus) dogs (C. prairie tailed northern and where of their range extent ludovicianus) dog (C. prairie black-tailed four near the city together of grouped are towns The 2004). (Seglund et al. Wyoming Cody, prairie black-tailed sympatric these of origin within the white-tailed isolated dogs,

INTRODUCTION INTRODUCTION 2 Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming Hoogland 2001).Female harassment may the first year (Clark and Stromberg 1987; may belessthan60percentand survival in year withanaverage of3.08to 3.88pups, 1979). Prairie dogsproduce onelitter per towns may beclumped(Hillmanetal. towns are not randomly distributed and prairie dogs(Hoogland 1996).Prairie dog different andblack-tailed for white-tailed regulated by theirnatural history, whichis may be Prairie dogdistributionandactivity distribution andabundance. many that factors prairie may dog affect however, thesetrends donotidentify the may helpidentify trends inpopulations; replicating historical surveys. These data ment ofbiases, limitations, anderrors when comparisons possiblewithacknowledge similar to thosein1980–1989,whichmade of habitat associations. Methodswere used 2001–2005 data were alsousedfor analysis indices ofdistribution andabundance. The BLM andFWS,summarizes changesin by conducted the the 1980–1989surveys byconducted theBLMin2001–2005with This compares report data from thesurvey 1985). Roberts 1:100,000 mapsin1988(Coy and U.S. Fish and Wildlife (FWS)on Service which was mappedindependently by the excluded theMeeteetse Metapopulation, Plan (Coy 1985). andRoberts Their study Prairie Ecosystem Dog Habitat Management for black-footed ferrets, andto produce a dog towns, better identify potential habitat These data were usedto inventory prairie communications, andground observations. photography, helicopter flights, personal including, butnotlimited to, infrared aerial prairie dogcoloniesmapping white-tailed oftechniquesa variety for locating and betweenconducted 1980and1989using for prairie were dogs.begin surveys Surveys Management’s (BLM)Cody Resource Area to area -

needed. studieshave Other new observed information is forBasin theBighorn characteristics appears to belimited and dog selection of certain soil and precipitation Weltzin etal. 1997).Data indicating prairie and Clark 1981;Coppock 1986; andDetling and prairie dogcontrol (Cambell efforts III be explained by many factors, including Habitat selection can vary greatly and might potential habitat might bemore successful. better defined,to conserve thenefforts defined. criteria Ifhabitat selection were based oncriteria that have notbeenfully establish newtowns, habitat theyselect was 2.4km. prairie As dogsemigrate and dog towns andtheirnearest neighbors mean distance between black-tailed prairie Hillman etal. that (1979)reported the and April andbetweenAugust. July 2.7 km duringemigration between March recorded prairie dogsmoving white-tailed emigrating nearly6.5km. Clark (1973) black-tailed(1958) observed prairie dogs dispersal distance of2.4km, butKoford anaveragereport black-tailed prairie dog observations. Garrett andFranklin (1988) and reported distances are usually maximum There have beenfew studiesondispersal fluctuations(Tileston and Lechleitner 1966). to berelated to prairie dogpopulation Immigration andemigration appear factors ofotheradults(Kofordthe animosity 1958). leaving theyoung behind, possiblydueto (Clark 1973). Some adults may also emigrate, occupy old, uninhabited burrow systems 1973). Young, dispersingmalestended to greatest (late (Clark andearlyJuly) June wastime whenthepopulation density Wyoming, pupsbeganto disperseat the and Franklin 1988).InAlbany County, in the coterie, encouraging dispersal (Garrett be caused by a shortage of unrelated females environmental conditions, vegetation barriers,

composition (Osborn andAllan 1949; vegetation seeddispersalandalter species Absence ofprairie increase dogscan woody 1973; Hoogland 1996; Weltzin etal. 1997). of tallervegetation thanblack-tailed (Clark prairie dogsarewhite-tailed more tolerant vegetation suchasgreasewood; however, ment berestricted can by insomecases tall and Lechleitner 1966).Prairie dogmove may eat more grasses andsedges(Tileston grasses, prairie dogs whilethewhite-tailed tailed prairie dogsmay eat mostly forbs and and Lechleitner 1966).Generally, black- and theavailability ofvegetation (Tileston withthetimeofyearTheir food habitsvary 1990). 1945;Holland andDetling (Bond increasecan shootnitrogen concentrations successional stages, andtheirherbivory ofearly forbs andotherplants typical 1997). Prairie dogsfeed mainlyonannual dogs (Osborn andAllan 1949; Weltzin etal. alsoinfluencecan thedistributionofprairie The vegetative andstructure community dehydration (Clark 1973;Parker etal. 1975). of annualplant growth orexisting growth saline uplandscorrelated withthecessation burrows insoilsthat are denseclay-clayey- zone selection forzone theprairie dogpopulation. selection and alsoanalyzes andprecipitation soiltype population indices spanning12to 26 years, This describestrends report incourse scale methods for the1980–1989BLMsurvey. used in2001–2005that were similarto was definedandinventory methods were biotic andabioticattributes. The studyarea dependent onmany known andunknown arePrairie dogdistributionandactivity vegetation encroachment. woodlands, indicating aversion to woody by shrubs andabsent insuccessional towns were present in grasslands uninvaded prairie active dog III(2001)report Riper Weltzin etal. 1997).Rosenstock and Van -

Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming 3 White-tailed prairie dog prairie White-tailed

THE PRAIRIE DOG TOWNS within the TOWNS THE PRAIRIE DOG study area (Total Population) were divided were Population) (Total study area the Basin metapopulations: two into and the Meeteetse Metapopulation Basin Metapopulation The Metapopulation. and south of the Montana was area east of the Shoshone line, State Wyoming of the Bighorn west Forest, National Creek and north of Shell Forest, National The 1). (Figure River Greybull the and was area Metapopulation Meeteetse 120, west of Highway towns defined as Wood north and the River of the Greybull Forest east of the Shoshone National River, North 40.00” 44° 19’ and south of Latitude observed 1). An white-tailed prairie (Figure 1973) (Clark of 2.7 km dog dispersal distance the Meeteetse separate used to was the Basin from Metapopulation Metapopulation.

STUDY AREA STUDY STUDY AREA STUDY Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming Figure 1. The area study includesthe Total Population, whichisequalto theBasinMetapopulation plustheMeeteetse Metapopulation areas. survey Forest Shoshone 0 Nttlonal 11 24 M 8 1216 - • mome-ters c:J ~ ~ -- Highway Sooonda,y Me«Keet9e 8Min R~OMlh + 1:614 t.ttca~lldotl N . 000 •nd Mecapopulallon Rettrnenl Buteau FOt9$1 SOMc:e of of or Patk Montana Indian Redamelion Und of SeMCe Oefen&e Mtt'l.-ntl'll Alfalrs '

Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming 5 Prairie rattlesnake occupying a black-tailed prairie dog burrow. dog prairie black-tailed a occupying rattlesnake Prairie

Aerial photography and former survey and former photography Aerial dog prairie in locating assisted results observations, ground In addition to towns. increase to interviewed were local residents were Ridges and roads detection rates. the visual maximize to or walked driven method most effective The surveyed. area appeared towns dog finding prairie for elevated from be binocular searches to and Roberts 1985). Aircraft (Coy locations precluded were but effective surveys are identified, was Once a town expense. due to identified prairie dog towns were resurveyed. resurveyed. were towns dog prairie identified as a polygon recorded was its location using a global positioning system 3). GeoExplorer (Trimble walking by defined boundaries were Town or the closest burrow to the perimeter A with a GPS unit. data recording mound, new if its boundaries considered was town during a identified a town did not overlap Because it took multiple years to accumulate accumulate to years multiple took it Because not instantaneous the surveysdata, are active Results represented descriptions. during the surveys, found dog towns prairie been complete. but surveys not have may FWS survey The did not include the same the 1980–1989 BLM methods used by activity burrow town survey; therefore, Metapopulation in the Meeteetse levels the to or compared not analyzed were 2001–2005 survey. former the replicate In an effortto survey surveys with similar methods, were November. through April conducted from conducted on BLM, Bureau Surveys were and private (BOR), state, of Reclamation in potential granted was lands when access which included dog habitat, prairie not were that communities vegetation forests, fields, irrigated bottoms, in river previously All and subalpine areas. alpine,

THE 1980–1989 SURVEY was identified THE 1980–1989 SURVEY identified was major surveyas including two efforts, the 1980–1989 BLM survey and the data survey Metapopulation (mapped Meeteetse in 1988), the FWS on 1:100:000 mylar by area surveyed the study which together issues and did with similar land access the of Fork South the up lands include not town dog prairie Active River. Shoshone screen these maps were from locations in GIS calculated were and areas digitized 1980–1989 Map 9.1 ESRI 2005. Active Arc assigned a new consecutive were towns sorting number by through “ID” town These and inactive towns. unknown the original survey to linked numbers were in GIS and on data referenced numbers, Dog Surveysheets in a new Prairie Binder in Microsoft a database and added to then were data These 2003. Access Office a new survey the to conducted by compared 2001 and 2005. BLM between

METHODS METHODS 6 Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming were since notcollected theywould not a town andsothey hadbeeninactive area, itcould notbedetermined how long to wildlife stilloccupyingimportant the 2006). Although towns inactive are very forbeen inactive many years etal. (Biggins they may berecently ormay inactive have are present throughout thelandscape, and burrow density. prairie Inactive dogtowns town were used to estimate theinactive however, burrows inactive withinanactive not collected or analyzed during this survey; et al. 2006).Data for towns inactive were vocalizations, andfresh excavation (Biggins individuals,such as:observed fresh scat, any recent prairie dogsignsofactivity A town was considered ifthere active were finest resolution was considered a “town.” burrowsnumber ofactive mappedat the Metapopulationthe Basin (Figure 1).Any as the Meeteetse Metapopulation and the Total Population and analyzed separately Data were analyzed for towns active within incorporated into thedatabase. assigned anidentification numberand (Coy 1985);thetown was andRoberts then Ecosystem Habitat Management Plan to theCody Resource Area Prairie Dog Inventory Form was filledoutaccording per 0.405ha(1.0ac).APrairie Dog Town within categories of≤10or>10burrows burrow densities were estimated and placed al. 1993).For thestudy, andinactive active estimates for burrow et (Biggins density the edgesoftown to calibrate observer 30 mparallel to eachothercontinuing to andsouthdistributed every oriented north 1meterused to widetransects conduct towns,low density ameasuringwheelwas season onrelatively high,medium,and previous survey. data collection Onceevery

instance, atown was found to have occurrence listed inthemapunit. For into range site areas usingthepercent The resulting areas soiltype were divided Potential areas. were calculated and withintheActual Total areas soiltype andprecipitation areas (NRCS), inthelate 1970sandearly1980s. Natural Resources Conservation Service now Conservationthe and theSoil Service, by theBLM initial field work performed precipitation zone data that are digitized, County’s soilsurveys,whichalsoincluded overlaid ontop ofPark andBighorn andPotentialIn GIS,Actual areas were (Clark 1973). for selection based on dispersal observations this buffer incorporated potential habitat white-tailed prairie dogs and it was assumed were analyzed asPotential useareas for buffer area around useareas the Actual analyzed useareas. asActual A2.7km prairie dogtownwhite-tailed areas were “Potential” use. The 2001–2005active analysis were basedon “Actual” and andprecipitation type Soil zone selection neighboring town. produce themeandistance to thenearest Nearest distances were averaged to towns tonearest thenext town edge. prairie dog boundaries ofallwhite-tailed measured distances from theouter edge statistics wereSummary produced for measured usingGISArcMap 9.1ESRI2005. prairie dogtownswhite-tailed were Nearest town distances for all2001–2005 abundance. indicate current prairie dogdistributionor

(244.0 ac)were usedinshalesites, 123.4ha percentage and it was assumed that 98.7 ha Therefore, thearea by was proportioned and 10percent that is “not specified.” 40 percent shale, 50percent salineupland, which generally hasrange sites with 474AD,247.1 ha(610.0ac)ofsoiltype and range site area within2.7km ofthe level,activity precipitation zone area, burrow density, thetwo categories of andthetwo categoriesthe two surveys of to test for significant associations between chi-square test wastwo-dimensional used inMicrosoftconducted Office Excel 2003.A Descriptive data analyseswere primarily were notentirely complete. foroccur private andthat soilsurveys lands aretype usedequivalently asoften asthey method assumesallrange sites withinasoil Potential available area (2.7km buffer). This andthe between theActual performed chisquare test wasa two-dimensional (8–10 in),and25.4–35.6cm(10–14in)]; [12.7–22.9 cm(5–9in),20.3–25.4 the three precipitation zones, whichoverlap Using the five most frequent range sites and precipitation zone were alsocalculated. Area andpercent occurrence ofeach weretypes excluded from theanalysis. (61.0 ac) were notnamed. Unspecifiedsoil (305.0 ac)were insalineuplands, and24.7ha Computation 2007). (Lowry and in VassarStats: Web Site for Statistical were inMicrosoft Excel performed 2003 were rejected at α=0.05.Statistical tests site statistical analysis. Nullhypotheses forper hectare theprecipitation andrange area withintowns. The Frequency unitwas prairie dogtownwhite-tailed edgeandthe

Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming 7 White-tailed prairie dog prairie White-tailed There were 66 active white-tailed towns 66 active white-tailed towns were There ac) with a 2,011.5 ha (4,970.6 occupying of 0.08 ha (0.2 ac) and a minimum area The maximum of 572.1 ha (1,413.8 ac). (73.7 ac) with a 29.8 ha was mean area deviation. 75.6 ha (186.7 ac) standard dog prairie black-tailed were towns Four 77.4 ha (191.3 ac), an totaling towns of 8 percent. increase 5 active white-tailed towns were There 615.2 ha (1,520.1 ac) with a occupying of 1.0 ha (2.5 ac) and a minimum area The maximum of 572.1 ha (1,413.8 ac). 123.0 ha (304.0 ac) with was mean area deviation. a 251.3 ha (621.0 ac) standard The 2001–2005 survey results for the for 2001–2005 survey results The Population: Total the for 2001–2005 survey results The Metapopulation: Meeteetse There were 49 active white-tailed towns 49 active white-tailed towns were There ac) with 4,736.7 ha (11,704.6 occupying of 0.04 ha (0.1 ac) and a a minimum area The maximum of 1,607.4 ha (3,972.0 ac). (239.0 ac) with a 96.7 ha was mean area deviation. 242.6 ha (599.5 ac) standard 56 active white-tailed towns were There 2,294.7 ha (5,670.3 ac) with a occupying of 0.008 ha (0.02 ac) and minimum area The a maximum of 631.4 ha (1,560.1 ac). 40.8 ha (100.8 ac) with was mean area deviation. an 89.9 ha (222.2 ac) standard dog prairie black-tailed were towns Two 71.3 ha (176.4 ac). totaling towns The 1980–1989 survey results for the for 1980–1989 survey results The Metapopulation: Meeteetse the for 1980–1989 survey results The Basin Metapopulation:

There were 105 active white-tailed towns towns white-tailed active 105 were There occupying 7,031.8 ha (17,374.9 ac) occupying of 0.008 ha with a minimum area (0.02 ac) and a maximum of 1,607.4 ha 66.4 ha was mean area The (3,972.0 ac). (164.0 ac) with a 178.3 ha (440.5 ac) were towns Two deviation. standard totaling dog towns prairie black-tailed 71.4 ha (176.4 ac). The 1980–1989 survey results 1980–1989 survey results The Population: Total the for Population Demographics

RESULTS RESULTS 8 Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming 1980–1989 Meeteetse Metapopulation. mean town area was larger thanthe 2001–2005 Meeteetse Metapopulation 1980–1989 Total Population (Table 1). The meantowna smalleractive area thanthe Metapopulation and Total Population had and Figure 2). The 2001–2005Basin Metapopulationfor theBasin (Table 1 Meeteetse Metapopulation, and39percent for the Total Population, 87percent for the was 71percentand the2001–2005survey occurring between the1980–1989surveys The percent decrease inoccupied area for the2001–2005survey. occupied by black-tailed prairie dogs area and4percent ofthearea was 96 percent oftheoccupied prairie dog tailed prairie dog population also comprised and astandard error of235.7. The white- minimum of48m,maximum13,243 was 1,963m,witha 1980–1989 survey nearest distance between towns for the and astandard error of279.7. The mean minimum of60m,maximum9,057 was 2,013m,witha 2001–2005 survey townsbetween white-tailed for the The meannearest neighbordistance for Metapopulation: theBasin The results 2001–2005survey towns totaling 77.4ha(191.3ac). Four towns were black-tailed prairie dog a 38.4ha(94.9ac)standard deviation. mean area was 22.7ha(56.0ac)with a maximumof271.7ha(671.5ac). The a minimumarea of0.08ha(0.2ac)and occupying 1,396.4ha(3,450.5ac)with There were towns white-tailed 61active

Table 1.Attributes prairiedogtowns ofwhite-tailed surveyed during1980–1989and2001–2005. 1980 and2001. Meeteetse Metapopulation, andBasinMetapopulation duringthetwo surveys beginningin by prairiedogs forFigure occupied white-tailed the 2.ComparisonTotal ofhectares Population, Mean Town Area Occupied AreaOccupied Town Abundance MetapopulationBasin Mean Town Area Occupied AreaOccupied Town Abundance Meeteetse Metapopulation Mean Town Area Occupied AreaOccupied HECTARES Town Abundance Total Population § § 0 ~ I I ~ I I I I I I I I I ..- Total 8 Population 40.8 ha(100.8ac) 2,294.7 ha(5,670.3ac) 56 96.7 ha(239.0ac) 4,736.7 ha(11,704.6ac) 49 66.4 ha(164.0ac) 7,031.8 ha(17,374.9ac) 105 1980–1989 Survey

Total § Population

- Meeleetst Metapopulatlon 22.7 ha(56.0ac) 1,396.4 ha(3,450.5ac) 61 123.0 ha(304.0ac) 615.2 ha(1,520.1ac) 5 29.8 ha(73.7ac) 2,011.5 ha(4,970.6ac) I 66

~ 2001–2005 Survey § ~ g Meeteetse - Met.ipopulation

- Ba~in 44 percent decrease 39 percent decrease 8 percent increase 79 percent increase 87 percent decrease 90 percent decrease 55 percent decrease 71 percent decrease 37 percent decrease I Metapopulation Percent Difference

,.., Basin § Met1population Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming 9

Inactive +71 percent +100 percent ila10 0£HSITY ≤10 >10 Table 2. Basin Metapopulation percent change change percent Metapopulation Basin 2. Table of surveys two the the time between during within active white active inactive and burrows as ≤ 10 or categorized dog towns tailed prairie per 0.405 ha (1.0 ac) categories. >10 burrows available were in clayey and shale range and shale range in clayey were available a higher frequency of illustrates This sites. - theoreti than the amount use of these sites them. Saline to uplands and cally available however, most common; were sites loamy have white-tailed dogs appear to prairie infrequent for a high selective propensity to appears There sites. range shale and clayey actual areas selected and the potential areas areas actual selected and the potential areas however, be some selection sites; loamy for the actual and potential loamy, shallow for 5). selection (Figure identical was percent -----.

BURROW 0 s1 4 ) 0 1 . . . 05 0 0 0 . 0.35 0.15 0 z => ~ ~ "" s - Figure 4. Basin Metapopulation frequency 4. Basin Metapopulation Figure of active and inactive within active white-tailed dog prairie density estimates burrow per 0.405 hectare as ≤10 or >10 burrows categorized towns the 2001–2005 survey. (1.0 ac) for ctiYe Ba i aacl!Ye a per 0.405 ha (1.0 ac) categories for the for (1.0 ac) categories per 0.405 ha significantly survey2001–2005 and were the with the time since associated 1980-1989 survey< 0.006, (X² = 12.76, P 2). Table 3 and 4; V = 0.24) (Figures Cramer’s tailed prairie dog towns, saline uplands dog towns, tailed prairie a major soil often were sites and loamy used. most frequently and were component and 22.8 made up 32.6 percent They active white-tailed of the total percent 5). (Figure respectively area dog town prairie be appear to sites Saline upland and loamy with white-tailed dog prairie associated dispersal potential the in soils as distribution, 5). (Figure less frequent relatively were area The two largest differences between the between differences largest two The >10

0£HSITY (Figure 5). (Figure

(X² = 77.67, BURROW 10 s 2 0 1 . 45 05 . . 0.4 0.3 0. 0 o.s 0.15 0.35 0.25 0 0 z => ~ ~ "" s - potential dispersal area (2.7 km) (2.7 km) dispersal area potential V = 0.017) P < 0.0001 and Cramer’s in activeOf white- the soil types present Habitat Associations Habitat A two-dimensional of the test chi-square a showed sites range most frequent five the actual between association significant selected white-tailed by areas site range potential to dogs when compared prairie within the available areas site dispersal Figure 3. Basin Metapopulation frequency 3. Basin Metapopulation Figure of active and inactive within active white-tailed dog prairie density estimates burrow per 0.405 hectare as ≤10 or >10 burrows categorized towns the 1980–1989 survey. (1.0 ac) for There was a trend towards more inactive more towards a trend was There the ≤10 and >10 burrows in both burrows Burrow density and activityBurrow within the have appears to Basin Metapopulation 2). Table 3 and 4; time (Figures changed over 10 Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming • • • • potentially occupied areas within 2.7 km.potentially occupied prairiedogtowns by white-tailed 25.4–35.6 cm(10–14in),occupied 2001–2005 active and Figure 6.Area oflandinprecipitation zones 12.7–22.9cm(5–9in),20.3–25.4(8–10and than what was available was in townthe active occurrence was greater Cramer’s V =0.047). The only zone where (X² =928.89,P<0.0001and prairie dogdistribution and white-tailed association between precipitation zones There alsoappearsto beasignificant within 2.7km. prairiedogtowns white-tailed active areas compared withpotentially occupied byFigure occupied 2001–2005 5.Area andpercent ofrangesite area actually %P«endal %P«endal %.lrtuol %Artuol P«ential P«ential Actual Actual He! = 41395 !! 518-6 (SU) 14.1 32.6 .... "g ::, ~ 115491.6 7345 5-9 55 40 34560 363.0 (Ly) 11.8 20.2 s ~ PREOPITATION 15695 160.9 (Sy) 10.1 .:>! -?;- ~ 9.1 RANGE 35857.1 610.7 8-10 34 17 site selection. prairiealso influence dog white-tailed town Cramer’s V values indicate othervariables Low for variability site selection. observed precipitation zones ofthe account for part zone (Figure 6). Thus, range sites and the 20.3–25.4cm(8–10in),precipitation SITE (Swly) 15641 - '.:!s 144.1 i 9.1 9.1 ZONE ~ 113.8 7555 (SIil ~ 4.4 7.8 18144.5 10-14 106 6 9 71.4 633 (Cy) 0 4.6 ~ .,,. £ .4

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 Cottontail rabbit occupying an inactive white-tailed prairie dog burrow. dog prairie white-tailed inactive an occupying rabbit Cottontail

urban areas, and other development and other development urban areas, the population. throughout interspersed there the Basin Metapopulation, Within between association also a significant was of active and inactive estimates categorical the 1980–1989 densities between burrow BLM surveys and the 2001–2005 survey. suitable for black-footed ferrets, although ferrets, black-footed suitable for where a suitable size to rebound it may where to be returned one day could ferrets found. was population source the recovery have appears to Basin Metapopulation The more, are There fragmented. more become less area which cover but smaller towns, both area and mean town area (occupied abundance while the town decreased, the Basin Within 1). Table - increased white-tailed dogs Metapopulation, prairie than the easily fragmented be more may highways, to due Metapopulation Meeteetse

study area, which is probably due to their due to which is probably study area, vegetation sagebrush for adaptations 1996). (Hoogland communities white-tailed two dog meta- prairie The Metapopulation (Meeteetse populations Total the within and Basin Metapopulation) changed in have appear to Population Meta- The Meeteetse ways. different ferrets black-footed where population, decline. the largest showed found, were covering towns, but larger fewer, are There in occupied decrease (a large much less area and an increase abundance, and town area This 1). Table - area in the mean town survivingsuggests that white-tailed prairie areas, core to contracted have dogs may and in part epizootic possibly resulting from other along with many plague, enzootic and Anderson factors (Menkens cumulative 2001). Currently Williams and 1991; Cully much less has become this metapopulation

of white-tailed prairie dog towns have have of white-tailed dog towns prairie large showing as a whole, decreased town categories: in four decreases mean town area, occupied abundance, 1 density (Tables and active burrow area, 2, 3, and 4). In other words, and 2; Figures were active white-tailed dog towns prairie inactive. and more smaller, fewer, dog prairie the black-tailed Although be it may slightly, increased population and so rare they are since vulnerable range. of the species’ the rest from isolated far the by dogs are White-tailed prairie dog species in the prairie most common THERE APPEARS TO HAVE BEEN MANY BEEN MANY HAVE TO THERE APPEARS CHANGES IN PRAIRIE DOG ACTIVITY and short time distribution during the relatively the between range) period (12–26 year black- the study area, Within surveys. two have appear to dog towns tailed prairie Population Total the while slightly, increased

DISCUSSION DISCUSSION 12 Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming historical surveys wherehistorical surveys thegoalisto difficult to accurately and precisely replicate limitations ofthestudydesign.Itisquite duetocautiously potential biasesand These results shouldbeinterpreted of1973. Act Species justification for listing under the Endangered declinecould leadto additional short-term prairie dogrange anddistribution.Another reduce the overallfurther white-tailed decreasesbefore occur, further whichcould reverse these trends should be implemented (Lomolino to 2001).Efforts andSmith in population distributionandabundance appears to have exhibited negative trends population withinthisstudyarea burrows, prairie dog thewhite-tailed from more burrows active to more inactive in occupied area and a proportional shift Basin. Thus, considering the overall decrease Bighorn population trends inthenorthern best available indicators ofprairie dog Trends inthesecolony attributes are the per.Biggins comm.). from 1988–2008(Dean Survey Geological the Meeteetse Metapopulation by theU.S. yearhas beendetected every sampledin be aplausibleexplanation since plague et al. 2004);however, enzootic plaguemay and Williams 200;Pauli etal. 2006;Seglund of theidentifiedcauses fordeclines (Cully declines were notanalyzed andmay beany demographic andtheotherobserved shift (Figures 3and4).Causal ofthis factors burrowin inactive estimates density estimates burrows ofactive andanincrease a shift from relatively higher to lower density there must have been certain factors causing during thetimebetween thetwo surveys, same methods. These results suggestthat databecause were withthe notcollected be madeintheMeeteetse Metapopulation Comparisons ofburrow could not density

burrow density, theerror was reduced by Although there are errors withestimating etal. studies)(Biggins large-scale 2006). mark-recapture, andmark-resight on rigorous methods (e.g., visualcounts, value andmay be impractical to use more methods haslong-term trend analysis replicate by usingsimilar ahistorical survey Lomolino 2001).Attempting and Smith to the studyarea etal. (Biggins 1993,2006; activity, occupied area, anddistributionfor be appropriate for trend estimates on resources were limited. The results should replicate theformer studyandtime used for thisstudywere designedto (Severson and Plumb1998),themethods more accurate populations indices to actual Although visualcounts might provide previous surveys. accurate mapsfrom thanhand-drawn the to maptowns, whichmay have beenmore ownership patterns. GPSunitswere used ofchangingland because two surveys however, may have changedbetween the were assumedto besimilar. Access to land, replicated andlimitations for bothsurveys methodswerethe 1980–1989surveys data tosurvey thepresent data, survey In order to compare 12–26year old the surveys. dog towns that were notdetected during possible inallcases, there may beprairie accessBecause to private landwas not administered lands, andlarger towns. prairie dogtowns onFederal andState- are likely underestimated, biasedtowards for bothsurveys. The results ofbothsurveys that thelevels were ofdetection thesame were includedinthisanalysis, orat least assumes that alltowns inthestudyarea etal. (Seglund 2004). activity This study inventory prairie dogdistributionand

a highrating oractive of>10inactive estimates fallwithinalow rating of≤10or using categories large enoughwhere habitat fragmentation (Clark 1973;Cambell quality, individualdispersalbehavior, and be related to many includinghabitat factors dog town spatial distribution,whichmay prairiedistance only describeswhite-tailed (1988) (2.4km). The meanminimum (1973) (2.7km) andGarrett andFranklin though similartoofClark observations measured between towns (2km) was less, The meanminimumdistance that was prairieby dogs(Clark white-tailed 1973). and precipitation zone habitat selection appropriate for comparisons ofrange site for apotential dispersaldistance andseems of 2.7km from allsurveyedtowns was used town. An area with aperimeter boundary precipitation, andproximity to thenearest distribution oftowns relative to soils, was aimedat describingthespatial The phaseofthisstudy habitat selection precise data. burrow andactivity density town providing for more accurate and have beendeployed prairie dog onevery and time, burrow could transects density frequency data. If there were more resources levelfor ascategorical estimating activity densities seems to be a reasonable approach to estimateobservers high andlow burrow burrows (1.0ac).Calibrating per0.405hectare type and local habitat andlocal variation.type Future adjusted dependinguponthedisturbance around prairie dogtowns may needto be and prairie dogtown connectivity. Buffers towns may helpmaintain current dispersal within a2km buffer zone from prairie dog managing for possibletown expansion population, minimizing disturbances and Considering declinesinthis theobserved 1986; Weltzin etal. 1997). III andClark 1981;Coppock andDetling

Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming 13

by anthropogenic factors should anthropogenic by

be studied further and may reveal additive additive be studied further reveal and may effects fluctuations. on normal population bacteria and recreational shooting can bacteria and recreational town affectsubstantially dog prairie 1968; Pauli et al. (Lechleitner persistence 2007) and it and Buskirk 2006; Pauli et al. plague has had overwhelming that is likely dynamics population dog prairie on influence Investigations comm.). (Dean Biggins per. selection habitat is natural on how influenced loamy sites, possibly those which are are possibly those which sites, loamy tracts agricultural primarily (Garrett now much how say to is hard 1982). It et al. and uses prohibit conditions land present their range. expanding dogs from prairie and practices such as grazing However, on the most occur regularly agriculture prairie displaced soils and have productive White-tailed 1958). (Koford dog towns whose soils, occupy dogs currently prairie and lower lack of depth, texture, alkalinity, converted not generally are precipitation sites less productive These agriculture. to history all life provide not adequately may be all the and perhaps may requirements fairly in a occupy is left to suitable land that or seasonally undisturbed undeveloped survey shows This 1982). et al. (Garrett state soil selection not represent and may current selection. habitat undisturbed historical, settlement before locations of town Studies were sites which range illustrate would used in a less disturbed ecosystem; be may evidence historical however, fast. eroding and is likely uncommon an have to appear Soils and precipitation effect selection dogs. by prairie on habitat areas these preferred from Displacement the observed to decline contribute may surely are There within the study area. variables and anthropogenic other habitat Plague dog decline. influencing the prairie flea-transmitted caused by epizootics

active area than what is proportionally than what active area km within the 2.7 available potentially that This suggests 5 and 6). (Figures buffer be more white-tailed dogs may prairie and site within this range successful range available Also, zone. precipitation white-tailed for less suitable prairie sites and outcrops such as rock dog towns, from nearly absent impervious were clays, the actual be limiting soils used and could et al. suitabilitydispersal and site (Garrett and precipitation site range The 1982). a analyses should be considered zone soil survey from scale assessment broad precipitation overlapping using information, et (Pauli collapse to be prone could matrix of 2006). Soils with high percentages al. which could lack permeability, would clay in water flooding and standing increase suggest and results study’s This burrows. a loamy that seem reasonable it would as tunnels be most suitable, soil would hold a proportion to of clay require may the addition of sand and silt yet the walls, easier burrowing. for allow would higher survivor - dogs can have prairie Since forbs, vigorous with more ship in areas they may historically and sedges, grasses, productive on more built their towns have zone categories, and may not reflect unique and may categories, zone these However, in localdiversity conditions. existing habitat refine help to may results suitability models. Soil white- types also be chosen by may their structural dogs for tailed prairie 1979). and Paulick (Flath components to tunnel in be difficult soils could Many soil may A shallow and inhibit dispersal. soils bedrock, enough depth to not allow of impervious be very would difficult clays or dig in, being directly in a wetland to of months could mean many floodplain other any lacking soils sandy and inundation,

through shooting or poisoning and through the to also be contributing loss may habitat these factorsobserved All should declines. effects the cumulative explain be studied to the observed to population contributing decline in the northern Bighorn Basin and their range. throughout loamy, upland, saline that study shows This shale range and loamy, shallow sandy, with associated significantly were sites selection site white-tailed dog town prairie within habitat available to when compared dogs White-tailed prairie 5). (Figure 2.7 km in the 12.7–22.9 cm the most area occupied and in the saline zone (5–9 in) precipitation and sites loamy however, site; range uplands the 20.3–25.4 cm (8–10 in) precipitation the highest proportion of actual have zone surveys could measure the mean minimum the mean surveys measure could for use it as an indicator again and distance be trending. may distribution how were sites dog town White-tailed prairie certain with range associated significantly study’s This zones. and precipitation sites with multiple an association showed results loamy, saline upland, including: sites range (1975) et al. Parker sites. and clayey shale, observed including dense similar sites upland selection. Otherclay-clayey-saline an effect on these have also likely variables V is relatively as the Cramer’s locations town 0.017). V = (Cramer’s low may selectionOther that habitat variables declines and distribution to be contributing vegetation condition, site range include: may health, soil permeability, and watershed of other or absence and the presence 1982; Ceballos et al. species (Garrett wildlife be may plague and Reading 1994). Sylvatic which frequently the number one factor, dogs in infected of prairie >99 percent kills Mortality 2001). Williams and (Cully colonies 14 Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming Endangered of1973,suchas: Act Species are BLMsensitive speciesorlisted underthe ecosystem andhabitat for species, which that provides aprey basefor acomplex aspecies for conserving trend isimportant Reversing negative the observed population for anecosystem dependent uponthe andincompleteThis specieslist isashort andSlobodchikoffregalis) 2000). (Bangert montanus), andferruginous hawk (Buteo cunicularia), mountain plover(Charadrius black-footed ferret, burrowing owl (Athene and Slobodchikoff 2000). andSavidge 1996;Bangert Desmond etal.Reading 1994;Miller 1994; (Campbell andClark 1981;Ceballos and presence anddistributionofprairie dogs

Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming 15 Black-tailed prairie dog, Brian Holmes, Colorado Division of Wildlife of Division Colorado Holmes, Brian dog, prairie Black-tailed Alteration of bison and black-tailed of bison and black-tailed Alteration by interaction dog grazing prairie Wildlife Journal of burning. prescribed 50(3):452-455. Management plan. management habitat ecosystem No. Management. of Land USDI Bureau WY-010-WHA-T14. plague Sylvatic comparisons of specific Journal of Mammalogy dogs. in prairie 82(4):894-905. owl influencing burrowing Factors (Speotyto nest densities and cunicularia) American Nebraska. numbers in western 136(1):143-148. Midland Naturalist characteristics of white-tailed dog prairie American Midland burrows. maternity 102(2):395-398. Naturalist Coppock, D.L. and J.K. Detling. 1986. and J.K. Detling. D.L. Coppock, dog Roberts. 1985. Prairie S.G. and D.A. Coy, - 2001. Inter Williams. and E.S. J.F. Cully, 1996. Savidge. and J.A. M.J. Desmond, 1979. Mound and R.K. Paulick. D.L. Flath, succession. Trans North American Wildlife Wildlife North American Trans succession. 10:229-234. Conference characteristics and vertebrate Colony of white-tailed and black- associates American Wyoming. dogs in tailed prairie 105(2):269-276. Midland Naturalist 1987. Houston. and B.R. S.C. Forrest, black-footed Wyoming habits of Food American Midland Naturalist ferrets. 117(1):208-210. and ethology of the white-tailed prairie : with a model of leucurus) dog (Cynomys dissertation, Ph.D. evolution. Cynomys Madison. Wisconsin, of University Press University Wyoming. in USA. Kansas, Lawrence, of Kansas. Campbell III, T.M. and T.W. Clark. 1981. T.W. and T.M. III, Campbell Clark, L. Richardson, T.W. T.M., III, Cambell 1973. A field study of the ecology T.W. Clark, 1987. and M.R. Stromberg. T.W. Clark, Bond, R.M. 1945. Range and plant and plant R.M. 1945. RangeBond, rodents

The Gunnison’s prairie dog structures a dog structures prairie Gunnison’s The high desert landscape as a grassland Journal of Arid engineer. keystone 46:357-369. Environments A. and R. Crete, A.H. Farmer, Oakleaf, B. evaluating for A technique 1993. Dood. Proceedings habitat. ferret black-footed on the Management of the Symposium the for Dog Complexes of Prairie of the Black-footed Reintroduction Service. Wildlife and USDI Fish Ferret. Biological Report 13:73-88. the abundance 2006. Estimating Ernst. (ed.). J.L., In: Hoogland, dogs. of prairie prairie of the black-tailed Conservation western Northdog: saving America’s Press. DC: Island Washington, grasslands. 94-107. p. Bangert, R.K. and C.N. Slobodchikoff. 2000. Bangert, R.K. and C.N. Slobodchikoff. L.R. Hanebury, Miller, B.J. D.E., Biggins, and A.E. Seery, D.B. Sidle, J.G. D.E., Biggins,

LITERATURE CITED LITERATURE LITERATURE CITED LITERATURE 16 Habitat Selection and Changes in the White-tailed and Black-tailed Prairie Dog Population within the Northern Bighorn Basin, Wyoming Koford, C.B. 1958.Prairie dogs, whitefaces, Knowles, C.J., C.J. Stoner, andS.P. Gieb. Hoogland, J.L. 2001. Black-tailed, Gunnison’s Hoogland, J.L. 1996.Cynomys ludovicianus. Holland, Detling. andJ.K. E.A., 1990. Hillman, C.N.,R.L.Linder, andR.B. Harlow, H.J. andMenkens Jr. G.E., 1986.A Garrett, M.G.,J.L. Hoogland, and W.L. Garrett, M.G.and W.L. Franklin. 1988. Lechleitner, M.I. R.R.,L.Kartman, 3:1-78. and bluegrama. Wildlife Monographs 84:71-74. dog towns by mountain plovers. Condor useofblack-tailed1982. Selective prairie Journal ofMammalogy82(4):917-927. and Utah prairie dogsreproduce slowly. 535:1-10. Mammalian Species 71(3):1040-1049. belowground nitrogen cycling. Ecology Plant response and to herbivory 102(1):185-187. ferrets. Naturalist AmericanMidland in areas inhabited by black-footed Dahlgren. 1979.Prairie dogdistribution Canadienne deZoologie 64(3):793-796. Canadian JournalofZoology/Revue dog, and Wyoming ground . tailed prairie dog, prairie white-tailed comparison ofhibernation intheblack- Naturalist 108(1):51-59 ludovicianus) . AmericanMidland black-tailed prairie dogs(Cynomys between anoldandanewcolony of Franklin. differences 1982.Demographic Mammology 69(2):236-250. black-tailed prairie dog. Journalof Behavioral ecology ofdispersalinthe central Colorado. Ecology 49(4):734-743. dogs (Cynomys insouth- gunnisoni) epizootic ofplagueinGunnison’s prairie Goldenberg, andB.W. Hudson.1968.An

Parker, J.L., G.L.Decker, L.Gray, andO. Osborn, B. andP. Allan. 1949. Vegetation of Miller, B., G.Ceballos, andR.Reading. 1994. Menkens, Jr. G.E., andS.H.Anderson. 1989. Menkens, Jr. G.E., andS.H.Anderson. 1991. Menkens, Jr. G.E., 1987. Temporal and Lowery, R.2007. Vassarstats: website for Lomolino, 2001. M.andG.A.Smith. Lehmer, andD.E. E.M. Biggins. 2005. 137 pp. of Agriculture, Conservation Soil Service. County area, Montana. U.S. Department Muller. ofCarbon survey 1975.Soil grass prairie. Ecology 30(3):322-332. town intall an abandonedprairie-dog Conservation 8(3):677-681. Biology The prairie dogandbioticdiversity. of Zoology 67:343-349. histories in Wyoming. Canadian Journal tailed prairie dogdemography andlife Temporal-spatial variation inwhite- 72(2):328-331. sylvatic plague. JournalofMammalogy prairie dogs during anepizootic of Population dynamicsofwhite-tailed Laramie, USA. University of dissertation, Wyoming, and life histories in Wyoming. Ph.D. dog (Cynomys leucurus) populations spatial variation prairie inwhite-tailed (accessed Feb. 25,2007). vassar.edu/lowry/VassarStats.html statistical computation. http://faculty. Mammalogy 82(4):937-945. the edgeoftheirrange. Journalof (Cynomys ludovicianus) towns near biogeographyDynamic ofprairie dog Journal ofMammalogy86:15-21. dogs across gradients ofelevation. ranging black-tailed andUtah prairie Variations intorpor patterns offree-

Weltzin, J.F., S.Archer, andR.K. Tileston, J.V. andR.R.Lechleitner. 1966. Plumb. andG.E. Severson, K.E. 1998. Seglund, Ernst, A.E. A.E., M.Grenier, B. Rosenstock, S.S.andC. Van III.2001. Riper Pauli, J.N. andS.W. 2007. Buskirk. Pauli, J.N., S.W. E.S. Buskirk, Williams, and 78(3):751-763. in atemperate savanna. Ecology regulation ofvegetation structure Heitschmidt. 1997.Small- Naturalist 75(2):292-316. central Colorado. AmericanMidland prairie dogsinnorth- and white-tailed comparisons of theblack-tailedSome 26(4):859-866. prairie dogs. Wildlife Bulletin Society population densitiesofblack-tailed Comparison ofmethodsto estimate assessment. UnpublishedReport. prairieWhite-tailed dogconservation Luce, A.Puchniak, andP. Schnurr. 2004. Management 54(3):226-232. woodland expansion. JournalofRange Breeding bird responses to juniper 71(1):103-108. chains. Journalof Wildlife Management for leadentering wildlife food a portal Recreational shootingofprairie dogs: 42(1):74-80. ludovicianus). Journalof Wildlife Diseases in theblack-tailed prairie dog(Cynomys W.H. Edwards. 2006.Aplagueepizootic 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. the Federal use by for or recommendation endorsement products does not constitute or commercial names, trade names, of company mention The