White-Tailed Deer Habitats in the Central Black Hills Author(S): Christopher S

Society for Range Management

White-Tailed Deer Habitats in the Central Black Hills Author(s): Christopher S. Deperno, Jonathan A. Jenks, Steven L. Griffin, Leslie A. Rice, Kenneth F. Higgins Reviewed work(s): Source: Journal of Range Management, Vol. 55, No. 3 (May, 2002), pp. 242-252 Published by: Allen Press and Society for Range Management Stable URL: http://www.jstor.org/stable/4003130 . Accessed: 11/01/2012 12:54

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp

JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

Allen Press and Society for Range Management are collaborating with JSTOR to digitize, preserve and extend access to Journal of Range Management.

http://www.jstor.org J. Range Manage. 55: 242-252 May2002 White-tailed deer habitats in the central Black Hills

CHRISTOPHER S. DEPERNO, JONATHAN A. JENKS, STEVEN L. GRIFFIN, LESLIE A. RICE, AND KENNETH F. HIGGINS

Authorsare Deer Project Leader, MinnesotaDepartment of Natural Resources, Farmland WildlifePopulations and Research Group,Madelia, Minn. 56062; Professor, Departmentof Wildlifeand Fisheries Sciences, South Dakota State University,Brookings, S.D. 57007; WildlifeBiologist, South Dakota Department of Game, Fish and Parks, 3305 W. South Street, Rapid City, S.D. 57702: RetiredSenior WildlifeBiologist, South Dakota Departmentof Game, Fish and Parks, 3305 W. South Street, Rapid City, S.D. 57702: Professor, South Dakota Cooperative Fish and Wildlife Research Unit, USGS-BRD, South Dakota State University,Brookings, S.D. 57007.

Abstract Resumen

White-tailed deer (Odocoileus virginianus dacotensis Zimm.) El numero de venados cola blanca (Odocoileus virginianus numbers in the central Black Hills have declined since the middle dacotensis Zimm.) del area central de Black Hills ha disminuido 1970s. Population status has been documented by a decline in desde mediados de la decada de los 70's. El estado de la hunter success, deer reproductive success, and fawn survival. poblacion ha sido documentado por una reduccion en el exito de Most management agencies believe habitat deterioration is the la caceria, la reproduccion exitosa del venado y la sobrevivencia primary cause of population decline in the Black Hills. We evalu- de los cervatos. La mayoria de las agencias de manejo creen que ated habitat selection for a white-tailed deer herd in the central el deterioro del habitat es la causa principal de la disminucion de la poblacion en Black Hill. Evaluamos la seleccion de habitat por Black Hills of South Dakota and Wyoming. From July 1993- un hato de venados cola blanca en la region central de Black Hill July 1996, 73 adult and yearling doe and 12 adult and yearling en South Dakota y Wyoming. De Julio de 1993 a Julio de 1996, a buck white-tailed deer were radiocollared and visually moni- 73 hembras adultas y juveniles y 12 machos adultos y juveniles tored. Habitat information was collected at 4,662 white-tailed de venado cola blanca se les coloco un radiotransmisor y se mon- deer locations and 1,087 random locations. During winter, white- itorearon visualmente. La informacion del habitat se colecto en tailed deer selected ponderosa pine- (Pinus ponderosa P. & C. 4,662 localidades de venado cola blanca y 1,078 localidades Lawson) deciduous and burned pine cover types. Overstory- aleatorias. Durante el invierno, el venado cola blanca seleccion6 understory habitats selected included pine/grass-forb, pine/bear- los tipos de cobertura de pino ponderosa (Pinus ponderosa P. & berry (Arctostaphylos uva-ursi (L.) Spreng.), pine/snowberry C. Lawson) deciduo y quemados. Los haibitats con cobertura (Symphoricarpos albus L.), burned pine/grass-forb, and superior/inferior seleccionados incluyeron pino/zacate-hierba, pino/ "Bearberry"(Arctostaphylos uva-ursi (L.) Spreng.), pino/ pine/shrub habitats. Structural stages selected included sapling- "Snowberry" (Symphoricarpos albus L.), pino quemado/zacate- pole pine stands with > 70% canopy cover, burned pine sapling- hierba y pino/arbusto. Las etapas estructurales seleccionados pole and saw-timber stands with < 40% canopy cover. During incluyeron poblaciones de plkantulasde pino con una cobertura summer, white-tailed deer selected pine-deciduous, aspen de copa mayor al 70%, poblaciones de plantulas de pino quema- (Populus tremuloides Michx.), aspen-coniferous, spruce (Picea do y poblaciones de pino aserrado con menos del 40% de cober- glauca (Moench) Voss), and spruce-deciduous cover types. tura. Durante el verano, el venado cola blanca selecciono cober- Overstory-understory habitats selected included pine/juniper turas del tipo pino-deciduo, alamo (Populus tremuloides Michx.), (Juniperus communis L.), aspen/shrubs, spruce/juniper, and ailamo-coniferas, picea (Picea glauca (Moench) Voss) y picea- spruce/shrub habitats. Structural stages selected included pine, deciduo. Los haibitats de cobertura alta-baja seleccionados aspen, and spruce sapling pole stands with all levels (0-40%, incluyeron haibitats de pino/enebro (Juniperus communis L.), ailamo/arbustos, picea/enebro y picea/arbustos. Los niveles 41-70%, 71-100%) of canopy cover. Results supported low habi- estructurales seleccionados incluyeron poblaciones de plantulas tat quality as a factor involved with the decline of the deer popu- de pino, ailamo y picea con todos los niveles (0-40%, 41-70%, lation. We recommend that habitat management techniques, 71-100%) de cobertura de copa. Los resultados apoyan la baja such as aspen regeneration and prescribed burns, be used to calidad del habitat como un factor involucrado en la disminucion improve the habitat base in the central Black Hills. de la poblaci6n de venados. Recomendamos que tecnicas de manejo del haibitat, tales como la regeneracion del alamo y los fuegos prescritos, sean utilizadas para mejorar el habitat base en la regi6n central de Black Hills.

This study was supported by Federal Aid to Wildlife Restoration Fund, Project W-75-R through the South Dakota Department of Game, Fish and Parks (Study Numbers 7563 and 7564). D. Flory, J. McCormick, D. Knowlton, S. Clark, B. Bol, D. Beck, and B. A. Hippensteel provided field assistance. We thank L. D. Flake, Key Words: aspen regeneration, Black Hills, habitat quality, M. K. Johnson, G. E. Larson, and J. Vandever for their review of this manuscript. habitat selection, Odocoileus virginianus dacotensis, prescribed T. Klinkner and J. Giudice provided editorial assistance. We wish to thank the burns, South Dakota, white-tailed deer, Wyoming USDA, Forest Service, Pactola and Harney Ranger Districts and all landowners that allowed access to their property throughout this study. Manuscript accepted 7 Sep. 01.

242 JOURNAL OF RANGE MANAGEMENT55(3) May 2002 White-taileddeer (Odocoileus virgini- type of cover used for fawningmay influ- southeast direction from high elevation anus dacotensis Zimm.) are an important ence the survivalrate of fawns, does seek summer ranges to low elevation winter economic resourcewithin the Black Hills isolation in areas where hiding cover is ranges and generally occurs between of South Dakota and throughout North abundant(King and Smith 1980, Fox and August and February (DePerno 1998, America. Williamson and Doster (1981) Krausman 1994). We hypothesized that Griffinet al. 1999). Typical springmigra- estimated that each white-tailed deer in white-taileddeer in the centralBlack Hills tion is in a northwestdirection from low NorthAmerica generates about $1,657 per select habitatson summerrange that con- elevation winter ranges to high elevation year throughconsumptive and non-contain abundant forage (stands with low summer ranges and generally occurs sumptiveuses. Restaurants,lodges, conve- overstorycover and significantshrub bio- between 17 and 23 May (DePerno 1998, nience stores, and gas stations receive mass) andcover (standswith high oversto- Griffinet al. 1999). Public land withinthe income from visitors attempting to ry cover and significant shrub biomass) studyarea is managedby the UnitedStates observe white-tailed deer (Martin and necessary to enhance reproductive suc- Departmentof AgricultureForest Service, Gum 1978, Wallace et al. 1991). cess, while selecting habitats on winter within the Pactola, Harney, and Elk Annually,the Black Hills deer herdgener- range that contain abundantforage and MountainRanger Districts, primarilyfor ates over $2 million dollars,in additionto cover necessaryto reducethermal stress. timberproduction and livestockgrazing (1 license fees (Richardson and Peterson June-31 October). 1974), with each residenthunter spending Winter range consists primarily of approximately$372/year (United States Study Area monotypic stands of ponderosa pine Fish andWildlife Service 1993). (Pinus ponderosa P. & C. Lawson) inter- White-taileddeer numbersin the central The Black Hills is an isolatedmountain- spersedwith standsof burnedpine, quak- Black Hills have declinedsince the middle ous area in western South Dakota and ing aspen (Populus tremuloides Michx.), 1970's (Griffin et al. 1992, Griffin 1994) northeastWyoming that extends approxi- and paper birch (Betula papyrifera andDePemo (1998) estimatedthis popula- mately 190 km northto south and 95 km Marsh.) (McIntosh 1949, Orr 1959, tion declined by 10-15% per year from east to west (Petersen 1993-1996. Population status has been 1984). Elevation of the documentedby a decline in huntersuccess Black Hills ranges from (McPhillips and Rice 1991), deer repro- 973-2,202 m above ductive success (Rice 1984, Hauk 1987, mean sea level (Orr McPhillips1990), and fawn survival(Rice 1959, Turner 1974). 1979). Additionally,because reproductive Annual mean tempera- and recruitmentrates have not increased turesare typicalof a con- with herd reductions,it appearsthat deer tinental climate and carryingcapacity in the centralBlack Hills range from 5-9? C with has declined(DePemo et al. 2000). Griffin extremes of -40-44? C et al. (1992) reportedthat most manage- (Thilenius 1972). Mean mentagencies believe the low productivity annual precipitation affectingthe centralBlack Hills deer herd ranges from 45-66 cm was directly related to long-term habitat (Orr 1959) and yearly : :. ~~~~~~~~~~~~~~~~~I deterioration. snowfallmay exceed 254 I,..- ~ ~ ~ ~ SUR We evaluatedhabitat selection by white- cm at higher elevations tailed deer in the central Black Hills of (Thilenius1972). South Dakotaand Wyoming.Habitats are The centralBlack Hills differentiatedby tree size and amountof study area (430 52' N to overstory cover present; characteristics 440 15' N-1040 07' W to that representage and structuralclass of 1030 22' W) includes vegetation within forested areas (Smith Penningtonand Lawrence 1962). Documentationof habitat use by countiesof SouthDakota bucks and does duringwinter and summer and Crook and Weston is necessary to develop area-specific counties of Wyoming strategies for maintaining deer popula- (Fig. 1). The study area tions. For example, in the northernBlack is composed of separate Hills, many habitatsused by white-tailed winter and summer deer differ from their relative availability rangesused by migratory on summer and winter ranges (Kennedy white-tailed deer 1992). Duringwinter, Moen (1968, 1976) (DePemo 1998, DePerno concluded that dense pine stands are et al. 2000, Griffinet al. importantto deer; these stands minimize 1995, 1999). In the cen- energy expendituresfor thermoregulation tral Black Hills, typical Fig. 1. Location of winter and summer ranges of white-tailed by reducing windchill and radiant heat autumn migration for deer in the central Black Hills, South Dakota and Wyoming, loss (Parkerand Gillingham 1990). During white-taileddeer is in a 1993-1996. springand summer,because condition and

JOURNAL OF RANGE MANAGEMENT55(3) May 243 Richardsonand Petersen Thilenius 1972, Appendix A. Common and scientific names of trees and shrubs included in the 1974, Hoffman and Alexander 1987). 'other shrubs' category. Primaryunderstory vegetation on winter range is characterized by snowberry Bearberry Arctostaphylosuva-ursi ((L.) Spreng.) (Symphoricarpos albus L.), spiraea Cherryspecies Prunusspp. (Spiraea betulifolia Pallas), serviceberry Currants Ribesspp. (Amelanchier alnifolia (Nutt.) Nutt. ex. M. Fleshyhawthorn Crataegussucculenta (Shrad. ex Link) Roemer), woods rose (Rosa woodsii Juniper Juniperuscommunis (L.) Lindl.), bearberry (Arctostaphylos uva- Leadplant Amorphacanescens (Pursh) ursi (L.) Spreng.), and cherry species Mountainbalm Ceanothusvelutinus (Dougl. ex Hook.) (Prunusspp.). Summerrange consists pri- Mountainmeadowsweet Spiraeabetulifolia (Pallas) marilyof ponderosapine and white spruce Oregongrape Berberisrepens (Lindl.) (Picea glauca (Moench) Voss) inter- Paperbirch Betulapapyrifera (Marsh.) spersed with small stands of quaking Ponderosapine Pinusponderosa (P. & C. Lawson) aspen (McIntosh 1949, Orr 1959, Quakingaspen Populustremuloides (Michx.) Thilenius 1972, Richardsonand Petersen Red raspberry Rubusidaeus (L.) 1974, Hoffman and Alexander 1987). Russetbuffaloberry Shepherdiacanadensis ((L.) Nutt.) Understoryvegetation on summerrange is Serviceberry Amelanchieralnifolia ((Nutt.) Nutt. ex M. Roemer) characterizedby Oregon grape (Berberis Snowberry Symphoricarposalbus (L.) repens Lindl.), juniper (Juniperus commu- Wartleberry Vacciniumscoparium (Leib ex Coville) nis L.), bearberry,snowberry, spiraea, and Whitespruce Picea glauca ((Moench)Voss) serviceberry. Willow Salix spp. Woodsrose Rosa woodsii(Lindl.) Materials and Methods Yellow rose Potentillafruticosa((Pursh) A. Love)

Capture Methods for white-tailed deer. Within the central 400-m2, circular plots centered on each White-taileddeer were capturedduring Black Hills, steep hills, deep draws, and deer observationsite (providingthe loca- February and March 1993-1996 using long migrationdistances limited data col- tion of the radiocollareddeer was visually modified,single-gate Clover traps (Clover lection activities to diurnal,visual obser- determinedwithout disturbing the animal) 1956) baited with fresh alfalfa (Medicago vations of deer and preventedthe use of and,to obtaina measureof relativehabitat sativa L.) hay. Deer were capturedon 4 other techniques (e.g., triangulation)for availability,at computergenerated random trapsites locatednortheast, northwest, and obtaining radiolocations. Furthermore, locations (i.e., sampled throughout the west of Hill City, S.D., on the McVey because of the terrainand inaccessibility study area) (Marcum and Loftsgaarden Burn deer winter range (Griffin et al. of many areas, attempts at spotlighting 1980, Kennedy 1992). General informa- 1995). Adult and yearling doe (n = 73) radiocollareddeer to obtainnocturnal data tion recorded at each location included: and buck (n = 12) white-taileddeer were were inefficient and represented a bias UTM location(north and east), and domi- fitted with radiocollars (Telonics Inc., toward deer that were more accessible. nant overstorytree species along with the Mesa, Ariz.; Lotek Engineering, Inc. Activity (feeding and bedding)was deter- most prominentunderstory vegetation. If Ontario, Canada), ear-tagged, aged by mined by radio signal intensityand speed 2 2 tree species provided canopy cover, lower incisorwear, and released.Captured of the pulse intervals (Beier and the species that provided the largest fawn white-tailed deer were ear-tagged McCullough 1988, Hansen et al. 1992, amountof cover was recordedas the pri- and released (Griffin et al. 1995). Each Weckerly1993). Deer locationswere plot- mary forest species; remaining species radiocollarused in this study containeda ted on 7.5-minute USGS topographical were recordedas secondaryspecies. mercurytip switchthat enabled determina- maps (scale, 1:24,000) and assigned Additional habitat characteristics tion of head-up and head-down position Universal Transverse Mercator (UTM) recordedat each location included:over- based upon signal intensity and differing coordinates (Edwards 1969, Grubb and story canopy cover (%), basal area pulse intervals (Beier and McCullough Eakle 1988). (m2/ha),diameter at breastheight (DBH) 1988). (cm), habitat association (pine, spruce, From July 1993-July 1996, individual Habitat Selection aspen), and vegetation structuralstage. radiocollareddeer were visually located Because separate winter and summer Percentoverstory canopy cover was mea- from the ground1-3 times per week. Deer rangesare used by migratorywhite-tailed sured using a spherical densiometer were radiotrackedat differenttime periods deer (DePerno1998, DePernoet al. 2000, (Lemmon1956). Basal area,the cross sec- to maximizeobservations of diurnalactiv- Griffin et al. 1995, 1999), we stratified tional area of trees at breast height, was ities (Kernohanet al. 1996) and to obtain data accordingto seasonalelevation shifts determinedusing a 10-factorangle gauge adequate sample sizes without violating made by each individualeach year (Apps (Hovind and Reick 1970). Diameter at the assumption of independentobserva- et al. 2001) and classified each deer loca- breastheight of each tree includedin the tions (White and Garrott1990). Kernohan tion and the correspondinghabitat infor- basal area count was measured, 1.37 m et al. (1996) demonstratedno differences mationas either winteror summerrange. above the ground(Ford-Robertson 1971), between diurnal and 24-hour habitat use Habitat information was collected from to the nearestcentimeter using a diameter tape. Vegetation structural stage units,

244 JOURNAL OF RANGE MANAGEMENT55(3) May 2002 Table 1. Percent forest type availability and use by doe and buck white-tailed deer during winter and summer in the central Black Hills, South Dakota and Wyoming, 1993-1996.

Winter Summer ForestType Does Bucks Availability (90 % CI) Does Bucks Availability (90% CI) (n = 1319) (n = 146) (n = 533) (n = 1496) (n = l81) (n = 563) Pine 68.5 68.5 72.8 (67.6 - 77.4) 44.7 - 54.7 58.1 (52.7 - 63.2) Pine/Deciduous 14.3+ 10.1 8.4 (5.6- 11.8) 20.9 + 15.5 + 9.4 (6.5 - 12.9) Aspen 2.3 1.3 2.1 (0.8 - 4.1) 5.4 + 3.9 2.3 (1.0 - 4.3) Aspen/Coniferous 2.0 2.0 2.6 (1.2 - 4.8) 8.8 + 7.2 + 3.6 (1.8 - 5.9) Spruce 0.1- 0.0- 0.8 (0.1 -2.2) 13.2+ 11.6 8.9 (6.1 - 12.2) Spruce/Deciduous 0.0 - 0.0 - 0.6 (0.0 - 1.9) 5.2 + 6.6 + 1.1 (0.3 - 2.6) BurnedPine 6.4 + 16.8 + 1.1 (0.3 - 2.8) 0.1 0.0 0.0 (0.0 - 0.8) Meadow 6.4- 1.3- 11.6 (8.4-15.5) 1.8- 0.6 - 16.7 (12.9-20.9) A positive sign (+) indicates significant habitatselection and a negative sign (-) indicates significant habitatavoidance. Significant levels for 90% confidence intervals were determined using the Bonferronimethod (Neu et al. 1974, Byers et al. 1984). which followed the Black Hills National Forest inventory system, were based on dominant overstory tree species, percent overstory canopy cover, and average DBH (Buttery and Gillam 1983, Rumble and Anderson 1992, 1993). Dominant tree species included ponderosa pine, spruce, and aspen. Meadow and burned pine habitats also were included. Pine, burned pine, spruce, and aspen stands were categorized based on stand age (Buttery and Gillam 1983). Structural stage categories included: 1 = grass/forb - 0 cm DBH; 2 = shrub/sapling - < 12.7 cm DBH; 3 = pole- timber - 12.7 - 22.8 cm DBH, and 4 = saw-timber - > 22.8 cm DBH. Structural stage categories 3 and 4 were further sepa- rated by percent canopy cover into: A = 0 - 40%, B = 41-70%, and C = 71-100% (e.g., A3B represents an aspen pole-stand with 41-70% canopy cover) (Kennedy Fig. 2. Burned pine habitat on winter range in the central Black Hills, South Dakota and Wyoming, 1993-1996. 1992, DePerno 1998). The overstory-understory relationships for pine, spruce, and aspen associations were determined by combining overstory characteristics (i.e., dominant tree species present, percent overstory canopy cover, basal area, average DBH) with dominant understory species present. Dominant understory species were determined in 15, 1-im2 plots systematically spaced within the 400-m2 area surrounding and including plot center (Daubenmire 1959, Kennedy 1992). Percent ground cover of grass, forbs, and shrubs < 1 m in height was visually estimated for each plot using the midpoint method as described by Daubenmire (1959).

Analytical Methods Forest type, overstory-understory habitat, and structural stages were pooled across individuals, years, and activities by Fig. 3. Aspen habitat on summer range in the central Black Hills, South Dakota and season. Deer locations were pooled for all Wyoming, 1993-1996.

JOURNAL OF RANGE MANAGEMENT55(3) May 245 Results

BetweenJuly 1993 andJuly 1996, 4,089 and 573 radiolocationswere obtainedfor does and bucks, respectively.To obtain a measure of relative habitat availability, habitatinformation was collected at 1,087 computer generated random locations sampled throughout the study area. Excluded from analyses were data on 1 radiocollaredbuck that remainedon winter rangethroughout the year and 1 radiocollared doe that demonstratedan abnor- mal migration pattern (DePerno et al. 1997).

Forest Type Forest type availabilityvaried between

I--~~ ~ ~ ~~~~~~~~~~~~~~~~~~~------winter and summer ranges (x2 = 68.56, df = 7, P < 0.001), and use of forest types (Table 1) differedfor does andbucks com- Fig. 4. Aspen/coniferoushabitat on summer range in the central Black Hills, South Dakota and Wyoming, 1993-1996. pared to forest type availability during winter (x2= 66.59, df = 7, P <0.001, x2 = 72.03, df = 7, P < 0.001) and summer(X2 = 238.77, df = 7, P < 0.00 1, X2= 30.32, df =7,P<0.001). During winter, pine/deciduous and burnedpine forests (Fig. 2) were selected (P < 0.05), whereasspruce, spruce/deciduous, and meadows were avoided (P < r 0.05) by does; remaining habitats were used in proportionto their availabilities. Buck use of burnedpine forest was nearly 3-times that of does and buck use of pine/deciduoushabitat was less (P < 0.05) than does and not significantly different from its availability. During summer, pine/deciduous,aspen (Fig. 3), aspen/coniferous (Fig. 4), spruce,and spruce/deciduous habitats were selected (P < 0.05), whereas pine and meadow habitatswere avoided (P < 0.05) and burnedpine was used in proportionto its availability by Bucksselected < Fig. 5. Burned pine/litter habitat on winter range in the central Black Hills, South Dakota does. (P 0.05) pine/decid- and Wyoming, 1993-1996. uous, aspen/coniferous,and spruce/deciduous habitats,while avoiding (P < 0.05) years of the studybecause of high season- 1992, McCleanet al. 1998). Selection for meadows;remaining habitats were used in al site fidelity (Progulske and Baskett a particularcover type, overstory-under- proportionto their availabilities.Overall, 1958, Ozoga et al. 1982, Tierson et al. story habitat, or structural stage was deer selected forest types that comprised 1985, Kennedy 1992, Nelson 1995, defined as use significantly greater than only 10%of the winterrange and 25% of Griffin et al. 1999). Because sufficient availability,while avoidancewas defined the summerrange. observationsper animal(x = 52.96 ? 5.20) as use significantly less than availability were recorded(Alldredge and Ratti 1986, (Neu et al. 1974, Byers et al. 1984, Overstory/UnderstoryType 1992) and becausehabitat availability was Kennedy 1992). Significance levels for Availability of overstory-understory assumedto be equal for all radiocollared 90%confidence intervals were determined types differed (X2 = 58.89, df = 14, P < animals,a chi-squaretest of homogeneity using the Bonferronimethod (Neu et al. 0.001) between summerand winterrange. was used to determinedifferences between 1974, Byers et al. 1984). All analyses Overstory/understoryuse differed (Table expectedand observed distributions of for- were performed using SYSTAT 2) for does comparedto availabilityduring est type, overstory-understoryhabitat, and (Wilkinson1990). winter (X2= 116.16, df= 14, P < 0.001) structuralstage (Jelinski 1991, Kennedy

246 JOURNAL OF RANGE MANAGEMENT55(3) May 2002 Table 2. Percent overstory-understory availability and use by doe and buck white-tailed deer during winter and summer in the central Black Hills, South Dakota and Wyoming, 1993-1996.

Winter Summer Forest Type Does Bucks Availability (90 % CI) Does Bucks Availability (90% CI) (n = 1000) (n = 124) (n = 470) (n = 1387) (n = 177) (n = 466) Pine/Grass/Forb 45.3 +1 37.9 + 14.9 (10.7- 19.7) 22.5 - 32.8 37.1 (31.0-43.3) Pine/Bearberry 2.0 + 11.3 + 0.2 (0.0-1.6) 5.8 - 9.0 9.4 (6.1-13.6) Pine/Juniper 0.0 0.8 0.0 (0.0-1.1) 24.3 + 16.4 + 6.7 (3.8 - 10.3) Pine/Snowberry 10.1 + 1.6 + 0.2 (0.0- 1.6) 0.7 0.6 0.6 (0.0-2.4) BurnedPine/Grass/Forb 6.8 + 17.7 + 0.0 (0.0- 1.1) 0.1 0.0 0.0 (0.0- 1.1) BurnedPine/Litter 25.4 - 22.6 - 78.7 (73.0 - 83.5) 12.0 - 6.8 - 25.8 (20.3 - 31.5) Pine/OtherShrubs2 5.4 + 4.0 + 0.2 (0.0-1.6) 2.3 5.1 + 0.9 (0.1-2.7) Aspen/Grass/Forb 2.3 2.4 3.0 (1.2 - 5.7) 8.6 7.9 5.4 (2.9 - 8.7) Aspen/Litter 0.5 0.8 1.5 (0.3 - 3.6) 1.0 0.6 0.4 (0.0 - 2.0) Aspen/OtherShrubs2 2.1 0.8 1.1 (0.2 - 3.0) 3.8 + 2.8 + 0.4 (0.0 - 2.0) Spruce/Grass/Forb 0.1 0.0 0.0 (0.0 -1.1) 7.6 7.4 5.8 (3.2 - 9.3) Spruce/Juniper 0.0 0.0 0.0 (0.0- 1.1) 4.2 + 4.0 + 0.9 (0.1 -2.7) Spruce/Wartleberry 0.0 0.0 0.0 (0.0 -1. 1) 3.1 1.7 2.2 (0.7 - 4.6) Spruce/Litter 0.0 0.0 0.2 (0.0- 1.6) 2.2 1.7 - 4.3 (2.1 -7.4) Spruce/OtherShrubs 0.0 0.0 0.0 (0.0- 1.1) 2.0 + 3.4 + 0.2 (0.0- 1.6) IA positive (+) sign indicates significant habitatselection and a negative sign (-) indicates significant habitatavoidance. Significance levels for 90% confidence intervals was determined using the Bonferronimethod (Neu et al. 1974, Byers et al. 1984). 2See Appendix A for a list of other shrubs.

and summer (X2= 203.11, df = 14, P < bucksdid not differ(x2 = 10.39, df = 14.0, pine/grass/forb,and ponderosapine/other 0.001), and for bucks duringwinter (X2= P = 0.73) fromhabitat availability. shrubs (AppendixA) were selected (P < 65.31, df = 14, P < 0.001). During sum- During winter, pine/grass/forb, pine/- 0.05), whereas burnedpine/litter (Fig. 5) mer,overstory-understory habitats used by bearberry, pine/snowberry, burned was avoided(P < 0.05) by does and bucks;

Table 3. Percentstructural stage availabilityand use by doe and buck white-taileddeer duringsummer and winterin the centralBlack Hills, South Dakotaand Wyoming,1993-1996.

Winter Summer Habitat Does Bucks Availability (90% CI) Does Bucks Availability(90% CI) (n = 477) (n = 47) (n = 532) (n = 786) (n = 99) (n = 562) P3A 9.6 4.3 - 8.5 (5.3- 12.4) 7.3 10.1 + 5.9 (3.3 -9.2) P3B 15.9 36.2 + 13.7 (9.6-18.4) 10.8 13.1 + 7.7 (4.7-11.3) P3C 22.2 + 25.5 + 14.7 (10.5-19.5) 10.4 + 12.1 + 4.8 (2.5-7.9) P4A 10.9 10.6 9.2 (5.9- 13.3) 17.2 23.2 + 14.2 (10.2- 18.9) P4B 12.4 - 8.5 - 17.1 (12.6 -22.2) 12.9 - 9.1 - 22.1 (17.1 - 27.4) P4C 9.2 - 0.0 - 19.2 (14.4-24.4) 7.0 - 3.0 - 11.9 (8.2- 16.3) A2 0.6 0.0 - 0.8 (0.1 - 2.5) 1.3 0.0 0.2 (0.0 - 1.4) A3A 0.6 0.0 - 0.6 (0.0 - 2.2) 2.5 + 4.0 + 0.5 (0.0 - 2.1) A3B 0.6 0.0 - 1.1 (0.2-3.1) 4.6 + 2.0 1.4 (0.3-3.4) A3C 2.1 0.0 - 2.1 (0.7 - 4.4) 2.8 + 2.0 0.9 (0.0 - 2.6) A4A 0.4 4.3 + 0.2 (0.0- 1.5) 1.2 0.0 - 0.9 (0.0-2.6) A4B 0.4 0.0 - 0.2 (0.0- 1.5) 1.3 0.0 - 0.9 (0.0-2.6) A4C 0.0 0.0 - 0.4 (0.0 - 1.8) 0.6 2.0 0.5 (0.0 - 2.1) S3A 0.0 0.0 - 0.0 (0.0 - 1.1) 1.7 2.0 0.7 (0.1 - 2.4) S3B 0.0 0.0 - 0.0 (0.0- 1.1) 2.9 + 3.0 + 0.4 (0.0- 1.7) S3C 0.0 0.0 - 0.0 (0.0- 1.1) 3.7 + 7.1 + 0.4 (0.0- 1.7) S4A 0.0 0.0 - 0.0 (0.0-1. 1) 4.6 + 0.0 - 2.1 (0.7-4.5) S4B 0.0 0.0 - 0.0 (0.0- 1.1) 3.4 3.0 3.9 (1.9-6.8) S4C 0.0 0.0 - 0.0 (0.0- 1.1) 2.0 3.0 3.6 (1.6-6.3) MD 9.0 4.3 - 11.3 (7.8- 15.7) 1.9 - 1.0 - 17.1 (12.7-22.0) BP3A 3.4 + 2.1 1.1 (0.2-3.1) 0.0 0.0 0.0 (0.0- 1.0) BP3B 0.6 2.1 + 0.0 (0.0- 1.1) 0.0 0.0 0.0 (0.0- 1.0) BP3C 0.0 0.0 - 0.0 (0.0- 1.1) 0.0 0.0 0.0 (0.0 - 1.0) BP4A 1.7 + 2.1 + 0.0 (0.0- 1.1) 0.0 0.0 0.0 (0.0- 1.0) BP4B 0.2 0.0 - 0.0 (0.0- 1.1) 0.0 0.0 0.0 (0.0- 1.0) BP4C 0.0 0.0 - 0.0 (0.0- 1.1) 0.0 0.0 0.0 (0.0- 1.0) I P = Pine; A = Aspen; S = Spruce;MD - Meadows; BP = BurnedPine. Significancelevels for 90% confidence intervals was determined using the Bonferroni method (Neu et al. 1974,Byers et al. 1984). 2= < 2.5 cm DBH; 3= 12.7 - 22.9 cm DBH; 4= > 22.9 cm DBH. A = - 40% canopy cover; B = 41 - 70% canopy cover; C = 71 - 100%canopy cover. 2A positivesign (+) indicatessignificant habitat selection and a negativesign (-) indicates significant habitatavoidance.

JOURNAL OF RANGE MANAGEMENT55(3) May 247 Table 4. Percent availability and use of habitats with and without shrubs selected by doe and buck medium age with medium to heavy white-tailed deer during winter and summer in the central Black Hills, South Dakota and canopies. Most of the remaining time was Wyoming, 1993-1996. spent in stands of aspen and spruce (Fig. 8) regardless of canopy cover. Overall, Does Bucks deer selected structural stages that com- Winter Summer Winter Summer prised only 30% of the winter range and With Shrubs 38% of the summer range. Forest Type n 1319 1496 146 181 availability (%) 9.6 25.2 9.6 25.2 use (%) 20.8 53.5 26.9 44.8 Discussion Overstory/Understory n 1000 1387 124 177 Clearly, in the central Black Hills, deer availability (%) 15.5 9.0 15.5 9.0 use (%) 69.6 36.6 72.6 31.7 spent most of their time in and were pri- StructuralStage marily supported by pine forested range, n 477 786 47 99 which dominates the landscape. However, availability (%) 29.7 38.3 29.7 38.3 the structural stage classification of the use (%) 44.2 66.8 72.4 76.8 Black Hills National Forest Inventory Without Shrubs System does not lend itself to clearly Forest Type n 1319 1496 146 181 explaining deer/habitat relationships. availability (%) 11.6 74.8 11.6 74.8 Selection of specific habitats by deer was use (%) 6.4 46.5 1.3 55.3 much better explained by the availability Overstory/Understory of understory plant communities, which n 1000 1387 124 177 provides, thermal cover, escape cover, and availability (%) 78.7 76.6 78.7 76.6 use (%) 25.4 42.4 22.6 50.3 food. Therefore, we more closely exam- StructuralStage ined these relationships by comparing deer n 477 786 47 99 use of habitats in relation to the availability (%) 61.1 52.8 61.1 52.8 presence/absence of shrubs (Table 4). On use (%) 35.2 24.2 17.0 13.1 winter range, deer use of forested stands with shrubs was 1.5 to 4.7 times greater than the availability of those habitats. On remaining habitats were used in propor- and selected medium to older age trees summer range, the relative difference tions to their availabilities. During sum- with medium to heavy forest canopies. between habitats containing shrubs select- mer, pine/juniper (Fig. 6), aspen/other Does spent twice as much time as bucks in ed by deer compared to the availability of shrubs, spruce/juniper, and spruce/other meadows and both sexes spent small those habitats ranged from 1.8 to 4.1 times shrubs were selected (P < 0.05), whereas amounts of time in burned pine stands. greater than those habitats without shrubs. bumed pine/litter was avoided (P < 0.05) During summer, deer spent > 60% of their Because hunter success (McPhillips and by does and bucks; pine/grass/forb and time in pine stands, primarily in stands of pine/bearberry was avoided (P < 0.05) by does and spruce/litter was avoided by bucks. All other habitats were used in pro- portion to their availabilities. Regardless of season, deer of both sexes selected understory range similarly and used types that comprised only 15% of the landscape on winter range and 9% of the landscape on summer range.

Structural Stage ..i,Z Availability of structural stages differed ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-1 ! (X2= 49.70, df = 25, P < 0.001) between winter and summer ranges. During winter, structural stage use for does (X2= 3.86, df e >^ ': . ! . = 25, P = 1.00) and bucks = 21.26, df ..2m ...... (X2 .i'i".'; * '2. 2 . . 0;f;...... :i:i. = 25, P = 0.68) did not differ from habitat I A~~~~~~~~~~~~~~~~A availability (Table 3). During summer, structural stage use differed for does (X2= 49.63, df = 25, P < 0.001) and bucks (X2= 39.75, df = 25, P < 0.001) compared to habitat availability. During winter, deer of both sexes spent Fig. 6. Pine/juniper habitat on summer range in the central Black Hills, South Dakota and 1993-1996. > 80% of their time in pine stands (Fig. 7) Wyoming,

248 JOURNAL OF RANGE MANAGEMENT55(3) May 2002 pine stands were selected because of the thermoregulatorybenefits they provide (Kennedy 1992). As air temperature declines, heat loss from the animal'ssur- face increases due to convection (Moen 1968). Therefore,deer may be experienc- ing a physiological benefit by occupying habitats on winter range that contain understoryand overstoryvegetation; habitats with high thermalcover characteristics (Moen 1976, Parker and Gillingham 1990). In addition to thermoregulatory benefits, we believe deer are selecting areas that contain shrubs (i.e., bearberry, juniper,and snowberry)because they are importantsources of forage necessary to wintersurvival of deer in the centralBlack Hills (Hill 1946, Schneeweis et al. 1972, Schencket al. 1972). This is furthersup- portedbecause pine habitatsdominated by snowberry,juniper, and bearberryare relatively rarein this regionof the Black Hills Fig. 7. Pine habitat on winter range in the central Black Hills, South Dakota and Wyoming, but were sought out and selected by deer. 1993-1996. Additionally, habitats with shrubs were selected at levels 1.5 to 4.7 times greater Rice 1991), deer reproductive success of white-tailed deer were composed of than the availability of those habitats. (Rice 1984, Hauk 1987, McPhillips1990, approximately40% ponderosapine, 30% Results indicate that approximately80% Hippensteel 2000), and fawn survival grass, 20% shrub, and 5% forbs, which of the habitat in the central Black Hills (Rice 1979, Benzon 1998) have not supportsthe contentionthat poor quality does not contain shrubs and thus, is not increasedwith herd reductions,Griffin et habitat is responsible for the long term acceptablefor deer. al. (1992) reportedthat most management populationdecline of white-taileddeer in Burned pine forests were selected by agencies believe the long term population the centralBlack Hills. white-tailed deer during winter but not decline and low productivity affecting during summer (burnedhabitats are rare central Black Hills white-tailed deer are Winter Range on summerrange). Burning speeds organ- directlyrelated to habitatdeterioration. In During winter, pine habitats that con- ic matterdecomposition rates and releases the central Black Hills, Hippensteel tained a shrub component were selected nutrientsinto the environment,promoting (2000) determinedthat winterrange diets by white-tailed deer. We believe these the productionof forage higher in protein (Einarsen 1946, Swank 1956, Sieg and Severson 1996). Similarly, researchers have concluded that burning in an unthinnedponderosa pine stand removed the littercomponent, initially increased the nutrientvalue of the herbaceousvegeta- tion, and stimulated growth of grasses, forbs, and shrubs,thereby increasingthe quality of the vegetation (Krefting 1962, Pearsonet al. 1972, Harestadand Rochelle 1982). However,most largerburned areas within the central Black Hills are > 40 years post-burn. Browse production increases for 3-5 years following a burn before returningto pre-burnlevels (Lay 1957, Taber and Dasmann 1957, Pearson l~~ ~ ~ ~~~~~...... et al. 1972). Consequently, deer use of *...... l||l ... _ll...... |I| burned areas may have been detected

Fig Spuc haia on su me rag in th ceta Blc Hils Sot Daoan because forage species (e.g., bearberry, Wymig 1993 -1996 .l |l snowberry,and juniper)important to deer (Hill 1946, Schneeweis et al. 1972, Schencket al. 1972) are essentiallyabsent in the dominantunburned pine communities of the centralBlack Hills.

JOURNAL OF RANGE MANAGEMENT55(3) May 249 On winter range, does selected open are limitedon summerrange in the central cover and forage are abundant(King and burnedpine and pine deciduous habitats, Black Hills (DePemo 1998, DePemo et al. Smith 1980, DePerno1998). Interestingly, whereasbucks selected closed pine stands 2000). Furthermore,grass and forbs com- does were located feeding (37%) more and avoided meadows.Interestingly, does posed < 30% of groundcover and shrubs thanbucks (14%). Therefore, we postulate were located feeding (52%) and bedding composed< 21% of the availableground that in the central Black Hills, does (48%) similarproportions of time, where- cover. These data coupled with approxi- migratefrom winterto summerrange just as bucks were located bedding (64%) mately48% littercover indicatethat grass, prior to parturitionto give birth in areas more often than feeding (36%). forb, and shrub forages are lacking on thatprovide thermal cover, maximumfor- Additionally,doe home rangeswere about summerrange in the central Black Hills age characteristics,and concealmentcover twice as large on winter (202.9 ha) range (DePerno 1998, DePerno et al. 2000). for fawns, whereasbucks migrateto sites compared to summer (130.9 ha) range Additionally,tall shrub sapling densities with high qualityforage on summerrange (Griffin et al. 1999). Largerwinter home were nearlythree times lower in the central to maximizebody condition. ranges and increasedtime spent foraging Black Hills (113.03 + 321.07 stems/ha) by does comparedto bucksmay be a func- compared to the northern Black Hills (3246.84 + 164.87 stems/ha;Hippensteel Summary and Management tion of the does requirementfor greater Implications quantitiesof forage during winter gesta- 2000), which suggests the central Black Hills lack tion, whereas bucks have lower energy sufficientunderstory vegetation and escape cover that are importantfor demands during this period and spend Resultsof this studyindicate that decid- white-tailed deer (DePerno 1998). more time loafing. Although,white-tailed uous cover types and habitatswith under- Furthermore,habitats with shrubs were deer normallyreduce food intake during story vegetationwere importantto white- selected at levels- 1.8 to 4.1 times greater winter (Short et al. 1969, Schultz et al. tailed deer in the centralBlack Hills. Sieg thanthe availabilityof those habitats,sug- and Severson (1996) hypothesized that 1993), we believe that adequateforage is gestingthat much of the habitatin the cen- limited (Spalinger et al. 1993) in this white-tailed deer densities in the Black tral Black Hills does not contain shrubs Hills region. For example, on winter range, have been reduceddue to the regen- andis not acceptablehabitat for deer. erationof ponderosapine stands,preven- aspen standsrepresent < 5% of the avail- During summer, structuralstages with tion of natural fires, and elimination of able habitat,understory shrubs (i.e., bear- 71-100% canopy cover, were selected by man-madefires. Absence of fire has sub- berry,juniper, and snowberry)and forbs white-taileddeer. Dense overheadcanopy stantiallyincreased ponderosa pine while represent< 30%,and littercomprises 57% cover providesa cool environment,which hinderingnew growth with negative con- of the available cover ground (DePerno may allow deer to avoid heat stress sequences for habitatdiversity, wildlife, 1998, DePernoet al. 2000). Furthermore, (Bunnell et al. 1986, Hoffman and and livestock interests (Richardson and > 90% of the forest standslack significant Alexander 1987) and reduce cutaneous Petersen 1974, Sieg and Severson 1996). understory vegetation or tall shrub water loss (Parker and Robbins 1984). Reducingconiferous overstory vegetation saplings (DePerno 1998, DePerno et al. Additionally,deer selected the relatively in and aroundaspen standscould increase 2000). Poor quality habitat and limited open aspen and spruce stands,suggesting understoryvegetation and forage diversity, availabilityof deciduous and shrubhabi- thathabitats containing understory vegeta- which would enhance opportunities for tats (DePerno1998), high doe mortalityin tion were important.Deciduous habitats herbivores to encounter higher quality early spring (i.e., 53.2% of radiocollared and habitats with shrubs were likely plants or plant parts (Ffolliot and Clary doe mortalityoccurred in spring,DePerno selected because they are important 1982, McConnell and Smith 1970, 1998, DePernoet al. 2000), an adultrepro- sources of forage and provide horizontal Seversonand Uresk 1988).Furthermore, if ductiverate of 1.33 fawns/doe,and a win- cover for predator avoidance (DePerno areas are logged first and then burned, ter range diet of 40% ponderosa pine 1998). forbs and shrubswill increaseand sprout (Hippensteel2000) indicate that does are In the centralBlack Hills, springmigra- growthof some species (e.g., chokecherry, nutritionallystressed on winter range in tion of white-taileddeer from low eleva- snowberry,and willow; Wrightand Bailey the centralBlack Hills. tion winterranges to high elevation sum- 1982) may be doubled (Krefting 1962, mer ranges generally occurs between 17 Harestadand Rochelle 1982). We recom- and 23 May (DePerno1998, Griffinet al. Summer Range mendthat agenciesresponsible for habitat 1999). This is about 3 weeks priorto the Duringsummer, deer selecteddeciduous manipulation/managementin the central peak date of parturition, which occurs forests that containedshrubs. Hill (1946) Black Hills give considerationto habitat approximately11 June(Benzon 1998). On concluded that aspen and associated treatmentsor modifications of on-going summer range, does selected deciduous shrubs were importantand highly palat- management (e.g., aspen regeneration, habitats that provide horizontal cover able to white-taileddeer duringsummer in logging, prescribedburning) that benefit (DePerno 1998) and an abundantsupply the northernBlack Hills. Aspen areashave white-taileddeer by significantlyincreas- of foragefor fawning(Smith and LeCount been determinedto be importantfor both ing deciduoushabitats and understory veg- 1979, King and Smith 1980, Bowyer and escape cover and forage for white-tailed etationon both winterand summerranges Bleich 1984, Riley and Dood 1984, deer throughoutthe Black Hills (Kranz in the centralBlack Hills. Huegel et al. 1986, Loft et al. 1987, Fox 1971, Schneeweiset al. 1972, Schencket Elk and livestock interestsmay be con- and Krausman1994, Main and Coblentz al. 1972, Kranzand Linder 1973, Kennedy tributing to deer decline in the central 1996, Uresk et al. unpublished data). 1992, Stefanich 1995). However, habitats Black Hills (DePerno1998, DePernoet al. Horizontal cover is importantto fawns; dominated by deciduous cover (< 12%) 2000) through competition for forage does seek isolation in areas where hiding

250 JOURNAL OF RANGE MANAGEMENT55(3) May 2002 (Jenks et al. 1996, Hippensteel 2000), Byers, C. R., R. K. Steinhorst, and P. R. Grubb, T. G. and W. L. Eakle. 1988. habitatdisturbance (Loft et al. 1987), dis- Krausman. 1984. Clarification of a tech- Recording wildlife locations with the placement (Crawford 1984, Loft et al. nique for analysis of utilization-availability UniversalTransverse Mercator (UTM) Grid data.J. Wildl.Manage. 48:1050-1053. System. USDA For. Serv. Res. Pap. RM- 1987, Loft 1988, Kie at al. 1991), survival Clover, M. R. 1956. Single-gate deer trap. 483. RockyMt. For. andRange Exp. Sta., Ft. (Smith 1982), and length of time on graz- Calif.Fish andGame. 42:199-201. Collins,Colo.. 3pp. ing allotments. Low to moderate cattle Crawford, H. S. 1984. Habitatmanagement. Hansen, M. C., G. W. Garner, and S. G. regimesmay be beneficialto elk and deer Pages 629-646 in L. K. Halls, editor.White- Fancy. 1992. Comparisonof 3 methodsfor (Skovlin et al. 1968). However, high tailed deer: ecology and management. evaluatingactivity of Dall's sheep. J. Wildl. Stackpole,Harrisburg, Penns., USA. Manage.56:661-668. dietary overlap between deer and elk Daubenmire, R. F. 1959. A canopy-coverage Harestad, A. S., and J. A. Rochelle. 1982. (49%), high pine consumption by deer method of vegetationalanalysis. Northwest Old-growthforests and black-taileddeer on (Hippensteel 2000), and length of cattle Sci. 33:43-64. VancouverIsland. Trans. North Amer. Wildl. grazing (i.e., in the central Black Hills DePerno, C. S. 1998. Habitat selection of a Nat. Resour.Conf. 47:363-373. livestockgrazing occurs from 1 Juneto 31 decliningwhite-tailed deer herd in the central Hauk, R. W. 1987. Deer managementsurveys, 1987. Pittman-RobertsonRept. W-95-R-22. the cattle grazingregime Black Hills, South Dakota and Wyoming. October)suggest Ph.D. Diss., S. Dak. State Univ., Brookings, S. Dak. Dept. of Game, Fish and Parks, presently practiced in the Black Hills is S. Dak. USA. Pierre,S. Dak.40pp. excessive and incompatiblewith improv- DePerno, C. S., S. L. Griffin, J. A. Jenks, Hill, R. R. 1946. Palatabilityratings of Black ing the white-taileddeer herd in the cen- and L. A. Rice. 1997. Unusualmigration by Hills plants for white-taileddeer. J. Wildl. tralBlack Hills. We recommendthe Forest a white-tailed deer fawn. Prairie Nat. Manage.10:47-54. Hippensteel, B. A. 2000. Factors affecting Service re-evaluatetheir currentgrazing 29:93-97. DePerno, C. S., J. A. Jenks, S. L. Griffin, nutritionalcondition of white-taileddeer in allotment (e.g., number of cattle and and L. A. Rice. 2000. Female survivalrates the centralBlack Hills, South Dakota.M.S. length of time) system as it relates to in a declining white-taileddeer population. Thesis, S. Dak. State Univ., Brookings, S. availabilityof deciduous and shrubhabi- Wildl. Soc. Bull. 28:1030-1037. Dak., USA. tats for white-tailed deer in the central Edwards, R. L. 1969. Archaeologicaluse of Hoffman, G. R. and R. R. Alexander. 1987. of the Black Hills National Black Hills. the Universal Transverse Mercator grid. Forestvegetation AmericanAntiquity 34:180-182. Forest of South Dakota and Wyoming: a Einarsen, A. S. 1946. Crudeprotein determi- habitattype classification.USDA Res. Pap. nation of deer food as an applied manage- RM-276. Rocky Mt. For. and Range Exp. Literature Cited ment technique.Trans. North Amer. Wildl. Sta., FortCollins, Colo. 48pp. Conf. 11:309-312. Hovind, H. J. and C. E. Reick. 1970. Basal Ffolliot, P. F. and W. P. Clary. 1982. area and point sampling:interpretation and Alldredge, J. R. and J. T. Ratti. 1986. application.Wis. Dept. Nat. Res. Tech. Bull. Comparisonof some statistical techniques Understory-overstoryvegetation relationships:an annotatedbibliography. USDA For. 23. 52 pp. for analysis of resource selection. J. Wildl. Huegel, C. N., R. B. Dahlgren, and H. L. Manage.50:157-165. Serv. Gen. Tech. Rep. INT-136. Intermtn. Gladfelter. 1986. Bedsiteselection by white- Alldredge,J. R. and J. T. Ratti. 1992. Further For. and Range Exp. Sta., Ogden, Utah. tailed deer fawns in J. Wildl. comparisonof some statisticaltechniques for 33pp. Iowa. Manage. analysis of resource selection. J. Wildl. Ford-Robertson, F. C., Ed. 1971. Termin- 50:474-480. Manage.56:1-9. ology of forest science technology practice Jelinski, D. E. 1991. On the use of chi-square Apps, C. D., B. N. McLellan, T. A. Kinley, and products: English language version. analyses in studies of resource utilization. and J. P. Flaa. 2001. Scale-dependenthabi- MultilingualFor. Term. Ser. 1. Soc. Amer. Can.J. For.Res. 21:58-65. tat selection by mountaincaribou, Columbia For.,Washington D.C. 349pp. Jenks, J.A., D.M. Leslie, Jr., R.L. Loch- Mountains, British Columbia. J. Wildl. Fox, K. B., and P. R. Krausman. 1994. miller, M. A. Melchiors,and F. McCollum, Manage.65:65-77. Fawning habitat of desert mule deer. III. 1996. Competitionin sympatricwhite- taileddeer and cattlepopulations in southern Beier, P. and D. R. McCullough. 1988. Southwest.Nat. 39:269-275. pine forests of Oklahoma and Arkansas, Motion-sensitiveradio collars for estimating Griffin, S. L. 1994. Seasonalmovements and USA. Acta Theriologica41 :287B306 white-taileddeer activity.J. Wildl. Manage. of in home range white-taileddeer the central Kennedy, J. F. 1992. Habitat selection by 52:11-13. Black South Pittman- Benzon, T. A. 1998. Mortalityand habitatuse Hills, Dakota, 1993. female white-tailed deer in the northern of white-tailed deer fawns in the central RobertsonReport W-75-R-35. South Dakota Black Hills, South Dakota and Wyoming. Black Hills, South Dakota 1994-1998. Dept. of Game, Fish and Parks, Pierre, S. M.S. Thesis, South Dakota State Univ., Pittman-RobertsonGame Rep. W-75-R-34. Dak. 13pp. Brookings,S. Dak. 65pp. S. D. Dept. of Game,Fish and Parks,Pierre, Griffin, S. L., J. F. Kennedy, L. A. Rice, and Kernohan, B. J., J. A. Jenks, D. E. Naugle, S.D. 43pp. J. A. Jenks. 1992. Movements and habitat and J. J. Millspaugh.1996. Estimating24-h Bowyer, R. T. and V. C. Bleich. 1984. Effects use of white-taileddeer in the northernBlack habitatuse patternsof white-taileddeer from of cattle grazing on selected habitats of Hills, S. Dak., 1991. Pittman-Robertson diurnaluse. J. Environ.Manage. 48:299-303. southernmule deer. Calif. Fish and Game. Report W-75-R-33. South Dakota Dept. of Kie, J. G., C. J. Evans, E. R. Loft, and J. W. 70:240-247. Game,Fish andParks, Pierre, S. Dak. 21pp. Menke. 1991. Foraging behavior by mule Bunnell, F.L., K.L. Parker, L.L. Kremsater, Griffin, S. L., L. A. Rice, C. S. DePerno, and deer:the influenceof cattle grazing.J. Wild. and F.W. Hovey. 1986. Thermoregulation J. A. Jenks. 1999. Seasonalmovements and Manage.55:665-674. and thermal cover of deer and elk on home rangesof white-taileddeer in the cen- King, M. M. and H. D. Smith. 1980. VancouverIsland: problem analysis. British tralBlack Hills, SouthDakota and Wyoming, Differentialhabitat utilization by the sexes of Columbia Ministries of Environment and 1993-97. S. Dak. Dept. of Game, Fish and muledeer. Great Basin Nat. 40:273-281. Forests. IWIFR-28. Victoria, British Kranz, J. J. 1971. A comparisonof aspenand Columbia. Parks,Pittman-Robertson Game Rep. W-75- pine communities in the northern Black Buttery, R. F. and B. C. Gillam. 1983. Forest R-34, Pierre,S. Dak.,USA. Hills. ecosystems. pp 34-71 In: R. L. Hoover and Griffiln,S. L., C. S. DePerno, J. A. Jenks, L. M.S. Thesis,South Dakota State Univ., D.L. Wills, eds. Managingforested lands for A. Rice, and D. A. Flory. 1995. Capture Brookings,S. Dak. 52 pp. wildlife. Colo. Div. of Wildl. in cooperation success of white-tailed deer in the central Kranz, J. J. and R. L. Linder. 1973. Value of with USDA Forest Serv., Rocky Mountain Black Hills, South Dakota. S. Dak. Acad. Black Hills forest communitiesto deer and Region,Denver, Colo.. 459 pp. Sci. 74:71-76. cattle.J. RangeManage. 26:263-265.

JOURNAL OF RANGE MANAGEMENT55(3) May 251 Krefting, L. W. 1962. Use of silvicultural Parker, K. L. and M. P. Gillingham. 1990. Skovlin, J. M., P. J. Edgerton, and R. W. techniquesfor improvingdeer habitatin the Estimates of critical thermalenvironments Harris. 1968. The influence of cattle man- lake states.J. For.60:40-42. for mule deer.J. Range.Manage. 43:73-80. agementon deer and elk. Trans.of the North Lay, D. W. 1957. Browse quality and the Parker K. L., and C. T. Robbins. 1984. Amer. Wildl. and Nat. Res. Conf. effects of prescribedburning in southernpine Thermoregulationin mule deer and elk. Can. 33:169-181. forests.J. For.55:342-347. J. Zool. 62:1409-1422. Smith, D. M. 1962. The practice of silvicul- Lemmon, P. E. 1956. A sphericaldensiometer Pearson, H. A., J. R. Davis, and G. H. ture, 7th edition. John Wiley and Sons, Inc. for estimatingforest overstorydensity. For. Shubert. 1972. Effects of wildfireon timber New York,N.Y. 578 pp. Sci. 2:314-320. and forage productionin Arizona.J. Range Smith, W. P. 1982. Status and habitatuse of Loft, E. R. 1988. Habitatand spatialrelation- Manage.25:250-253. Columbian white-tailed deer in Douglas ships between mule deer and cattle in a Petersen, L.E. 1984. Northern Plains. pp. County,Oregon, Diss., Oregon State Univ., Sierra Nevada forest zone. Diss., Univ. of 441-448 In: L. K. Halls, ed. White-tailed Corvallis,Ore., USA. California,Davis, Calif.,USA. deer: ecology and management.Stackpole Smith, R. H. and A. LeCount. 1979. Some Loft, E. R., J. W. Menke, J. G. Kie, and R. Books, Harrisburg,Penn. 870pp. factorsaffecting survival of desertmule deer C. Bertram. 1987. Influenceof cattle stock- Progulske, D. R. and T. S. Baskett. 1958. fawns.J. Wildl.Manage. 43:657?665. ing rate on the structuralprofile of deer hid- Mobility of Missourideer and their harass- Spalinger, D. E., C. T. Robbins, and T. A. ing cover.J. Wildl.Manage. 51:655-664. mentby dogs. J. Wildl.Manage. 22:184-192. Hanley. 1993. Adaptive rumenfunction in Main, M. B. and B. E. Coblentz. 1996. Sexual Richardson,A. H., and L. E. Peterson. 1974. elk (Cervus elaphus nelsoni) and mule deer History and Managementof South Dakota (Odocoileus hemionus hemionus). Can. J. segregationin Rocky Mountainmule deer.J. Deer. South Dakota Dept. Game, Fish and Zool. 71:601-610. Wildl.Manage. 60:497-507. Parks Publ. Bull. No. 5. Pierre, S. Dak. Stefanich, M. R. 1995. Movementsand habitat Marcum, C. L. and D. 0. Loftsgaarden. 113pp. use of white-taileddeer in the northwestern 1980. A nonmappingtechnique for studying Riley, S. J. and A. R. Dood. 1984. Summer Black Hills of Wyomingand South Dakota. habitat preferences. J. Wildl. Manage. movements, home range, habitat use, and M.S. Thesis, Univ. of Wyo., Laramie,Wyo. 44:963-968. behavior of mule deer fawns. J. Wildl. 46 pp. Martin, W. E. and R. L. Gum. 1978. Manage.48:1302-1310. Swank, W. G. 1956. Proteinand phosphorous Economicvalue of hunting,fishing, and gen- Rice, L. A. 1979. Mortalityrates of fawn age contentof browse plants as an influenceon eral rural outdoor recreation. Wildl. Soc. class in South Dakota deer populations, southwesterndeer herd levels. Trans.North Bull. 6:3-7. 1978-1979. Pittman-RobertsonReport W- Amer.Wildl. Conf.21:141-158. McClean, S. A., M. A. Rumble, R. M. King, 75-R-21. S. Dak. DepartmentGame, Fish Taber, R. D. and R. F. Dasmann. 1957. The and W. L. Baker. 1998. Evaluation of andParks, Pierre, S. Dak. 17 pp. dynamicsof three naturalpopulations of the resource selection methods with different Rice, L. A. 1984. Fawnmortality rates in South deer Odocoileus hem ionus columbianus. definitionsof availability.J. Wildl. Manage. Dakota deer populations, 1977-1981. Ecol. 38:233-246. 62:793-801. Pittman-RobertsonReport W-75-R-26. S. Thilenius, J. F. 1972. Classification of deer McConnell, B. R., and J. G. Smith. 1970. Dak. Dept. Game, Fish and Parks,Pierre, S. habitat in the ponderosapine forest of the Response of understoryvegetation to pon- Dak.45pp. Black Hills, SouthDakota. USDA For. Serv. derosa pine thinningin easternWashington. Rumble, M. A. and S. H. Anderson. 1992. Res. Pap.RM-91 . FortCollins, Colo. 28 pp. J. RangeManage. 23:208-212. Stratificationof habitatsfor identifyinghabi- Tierson, W. C., G. F. Mattfield, R. W. Sage, McIntosh, A. C. 1949. A botanicalsurvey of tat selection by Merriam'sTurkeys. Great Jr., and D. F. Behrend. 1985. Seasonal the Black Hills of SouthDakota. Black Hills BasinNat. 52:139-144. movementsand home rangesof white-tailed Engin.28:1-74. Rumble, M. A. and S. H. Anderson. 1993. deer in the Adirondacks.J. Wildl. Manage. McPhillips, K. 1990. Deer managementsur- Habitat selection of Merriam's turkey 49:760-769. veys, 1989. Pittman-RobertsonReport W-95- (Meleagris gallopavo merriami) hens with Turner, R. W. 1974. Mammalsof the Black R-24. South Dakota Departmentof Game, poultsin the BlackHills, S. Dak. GreatBasin Hills of South Dakotaand Wyoming.Misc. Fish andParks, Pierre, S. Dak. 40pp. Nat. 53:131-136. Publ. Mus. Nat. Hist. Universityof Kansas. McPhillips, K. and L. A. Rice. 1991. Deer Schenck, T. E., III, R. L. Linder, and A. H. 178 pp. managementsurveys, 1991. South Dakota Richardson.1972. Food habitsof deerin the United States Fish and Wildlife Service. Department of Game, Fish and Parks, Black Hills, Part II: SouthernBlack Hills. 1993. National Survey of Fishing, Hunting, Pittman-RobertsonGame Report W-95-R-25, South Dakota Agr. Exp. Sta. Bull. 606, S. and Wildlife-Associated Recreation. U.S. Pierre,S. Dak. 49 pp. Dak. StateUniv., BrookingsS. Dak. 35pp. Govern. PrintingOff., WashingtonD.C. 41 Moen, A. N. 1968. Surface temperaturesand Schneeweis,J. C., K. E. Severson, and L. E. radiantheat loss PP. from white-tailed deer. J. Petersen. 1972. Food habits of deer in the Wallace, M. S., H. L. Stribling, and H. A. Wildl. Manage.32:338-344. BlackHills, Part I: NorthernBlack Hills. South Clonts. 1991. Effects of hunterexpenditure Moen, A. N. 1976. Energy conservation by DakotaAgr. Exp. Sta. Bull. 606, SouthDakota distributionon communityeconomics. Wildl. white-tailed deer in winter. Ecol. StateUniv., Brookings, S. Dak.35 pp. Soc. Bull. 19:7-14. 57:192-198. Schultz, S. R., M. K. Johnson, and A. E. Weckerly, F. W. 1993. Intersexualresource Nelson, B. E. 1995. Use of a Geographic Hindrichs. 1993. Effects of spring/summer partitioningin black-taileddeer: a test of the Informationsystem for determiningwhite- body mass gain and sex ratio on fall/winter body size hypothesis. J. Wildl. Manage. tailed deer habitatuse in the NorthernBlack body mass loss of adult, male white-tailed 57:475-494. Hills of South Dakota and Wyoming. M.S. deer.Small Rum. Res. 10:183-188. White, G. C. and R. A. Garrott. 1990. Thesis, SouthDakota State Univ., Brookings, Severson, K. E. and D. W. Uresk. 1988. Analysis of wildlife radio-tracking data. S. Dak. USA. Influenceof ponderosapine overstoryon for- AcademicPress, New York,N.Y. 383 pp. C. R. Neu, W., C. Byers, and J. M. Peek. age qualityin the Black Hills, SouthDakota. Wilkinson, L. 1990. Systat:the systemfor sta- 1974. A techniquefor analysisof utilization- GreatBas. Nat. 48:78-82. tistics.Systat, Inc. Evanston,Ill. 750 pp. data. availability J. Wildl. Manage. Short, H. L., J. D. Newsom, G. L. McCoy, Williamson, L. L. and G. L. Doster. 1981. 38:541-545. and J. F. Fowler. 1969. Effects of nutrition Socioeconomicaspects of white-taileddeer H. K. 1959. and streamflow Orr, Precipitation and climate on southerndeer. Trans.North diseases.pp 434-439 In: W.R. Davidson,ed. in the Black Hills. USDA Station Pap. 44. Amer.Wildl. Conf. 34:137-146. Diseases and parasitesof white-taileddeer. Rocky Mt. For. and RangeExp. Station,Fort Sieg, C. H. and K. E. Severson. 1996. Tall Timbersresearch Station, Tallahassee, Collins,Colo. 25pp. Managing habitatsfor white-tailed deer in Fla. 472 pp. Ozoga, J. J., L. J. Verme, and C. S. Bienz. the BlackHills andBear Lodge Mountains of Wright, H. A. and A. W. Bailey. 1982. Fire 1982. Parturitionbehavior and territoriality South Dakota and USDA For. John New in Wyoming. Ecology. Wiley & Sons, York, white-taileddeer: impact on neonatalmor- Serv. Gen. Tech. Rep. RM-GTR-274.Rocky N.Y. 501pp. tality.J. Wildl.Manage .46:1-11. Mt. For. and Range Exp. Station, Fort Collins,Colo. 24pp.

252 JOURNAL OF RANGE MANAGEMENT55(3) May 2002