Responses of Female White-tailed Deer Home-Ranges to
Increased Resource Availability
Shawn M. Crimmins, John W. Edwards, Tyler A. Campbell, W. Mark Ford, Patrick D. Keyser, Brad F. Miller, and Karl V. Miller
Northeastern Naturalist, Volume 22, Issue 2 (2015): 403–412
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2015 NORTHEASTERN NATURALIST 22(2):403–412
Responses of Female White-tailed Deer Home-Ranges to
Increased Resource Availability
Shawn M. Crimmins1, 2,*, John W. Edwards1, Tyler A. Campbell3, W. Mark Ford4,
Patrick D. Keyser5, Brad F. Miller6, and Karl V. Miller7
Abstract - Management strategies designed to reduce the negative impacts of overabundant
Odocoileus virginianus (White-tailed Deer) populations on forest regeneration may
be influenced by changes in both population density and timber harvest. However, there
is conflicting evidence as to how such changes in per capita resource availability influence
home-range patterns. We compared home-range patterns of 33 female White-tailed
Deer from a low-density population at a site with abundant browse to patterns of a sample
of >100 females prior to a 75% reduction in population density and a doubling in timber
harvest area. Home-range and core-area sizes were approximately 3 times larger than were
found prior to population decline and timber harvest increase, consistent with predictions
related to intraspecific competition. We also observed greater site fidelity than previously
exhibited, although this may be an artifact of increased home-range sizes. Our results
support previous research suggesting that White-tailed Deer home-range size is inversely
related to population density and is driven, in part, by intraspecific competition for resources.
Relationships among population density, resource availability, and home-range patterns
among female White-tailed Deer appear to be complex and context specific.
Introduction
Variability in the size of an animal’s home range, i.e., the area used during everyday
activities (Burt 1943), is a function of numerous biotic and abiotic factors.
Although intrinsic factors such as mating system may influence seasonal patterns
in home-range size (Clutton-Brock 1989), intraspecific competition for resources
directly affects home-range size (Burt 1943, Sanderson, 1966). In many species,
individual home-range size is inversely related to population density (Brown 1969,
Getz 1961). This relationship is generally explained as a function of habitat quality,
with high-density populations occurring in areas with more resources available and
thereby requiring less movement to meet energetic or other resource requirements.
Additionally, free distribution in habitat selection would also lead to an inverse
relationship between population density and home-range size due to territorial
1Division of Forestry and Natural Resources, West Virginia University, Morgantown, WV
26506. 2Current address - University of Wisconsin, Department of Forest and Wildlife
Ecology, Madison, WI 53706. 3East Wildlife Foundation, San Antonio, TX 78216. 4US Geological
Survey, Virginia Cooperative Fish and Wildlife Research Unit, Virginia Polytechnic
Institute and State University, Blacksburg, VA 24061. 5Department of Forestry, Wildlife,
and Fisheries, University of Tennessee, Knoxville, TN 37996. 6National Wild Turkey Federation,
Bristol, TN 37849. 7Warnell School of Forestry and Natural Resources, University
of Georgia, Athens, GA 30602. *Corresponding author - scrimmins@wisc.edu.
Manuscript Editor: Kurt Moseley
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interactions (Fretwell and Calver 1969). However, population density is not always
monotonically related to habitat quality such that areas with higher population
density equate to areas with higher habitat quality (VanHorne 1983). In highly
philopatric species, such as Odocoileus virginianus Zimmerman (White-tailed
Deer, hereafter Deer), it has been suggested that population density and home-range
size are directly related (Kilpatrick et al. 2001, Tiersen et al. 1985). These discrepancies
in hypothesized relationships make it difficult to predict the relationship
between population density and home-range patterns in any individual population.
Although the effects of population density on Deer recruitment (Keyser et al.
2005a), physical condition (Garroway and Broders 2005, Keyser et al. 2005b),
and herbivory (Miller et al. 2010), are well known and generally consistent, their
effects on space-use patterns are inconsistent, especially in forested landscapes.
For example, on coastal islands in South Carolina, seasonal home-range sizes of
suburban Deer increased in response to a 50% reduction in population density
(Henderson et al. 2000). Similarly, home-range sizes of O. virginianus clavium
Barbour and Allen (Florida Key Deer) decreased as population densities increased
(Lopez et al. 2005). Conversely, Deer in suburban Connecticut decreased their
annual home-range sizes immediately after the implementation of a herd reduction
program (Kilpatrick et al. 2001). Studies of female Deer herds in enclosures
have found an inverse relationship between population density and home-range
size (Williams and DeNicola 2002). In a forested landscape in the Adirondacks
of New York, McNulty et al. (1997) found that home-range size did not change in
response to a localized removal of 80% of the female Deer in their study population,
but did find that levels of philopatry decreased at moderate to relatively low
population density exhibited in their study area (6–12 Deer/km2). Populations
occurring at higher density or that have undergone a more substantial change in
density may exhibit different patterns. For example, populations occurring at
higher densities should experience greater levels of intraspecific competition
among individuals for finite resources than those at low density, and thus may be
expected to exhibit greater responses to reductions in population size. However,
McNulty et al. (1997) remains one of the only studies to address the effects of
population density on both home-range size and patterns of philopatry.
Population density can affect home-range patterns by changing the availability
of resources or through the amount and intensity of intraspecific interactions. An
increase in resource availability, as would occur after a reduction in population
size, should lead to a decrease in home-range size because animals can obtain
the required amount of resources over a smaller area (Kilpatrick et al. 2001).
Conversely, reducing population size may also decrease aggressive intraspecific
interactions, which could result in increasing home-range size (e.g., Henderson et
al. 2000). Because the effects of changes in resource availability on White-tailed
Deer home-range sizes are relatively unknown, we contrasted home-range sizes
and levels of philopatry in a population prior to and following a >75% reduction
in Deer density and an independent doubling in the amount of forage availability
Northeastern Naturalist Vol. 22, No. 2
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in a forested landscape of the Central Appalachians of West Virginia. Our
objectives were to determine how: (1) seasonal home-range and core-area sizes
and (2) inter-annual site fidelity changed in response to reduced population density
and increased timber harvest. We hypothesized that home-range and core-area
sizes would decrease due to a presumed increase in per-capita resource availability
resulting from both a reduction in population density and increase in timber
harvest, and that philopatry would increase due to a reduced need for extensive
movements to acquire resources.
Methods
Study area
We conducted our study in the MeadWestvaco Wildlife and Ecosystem Research
Forest (MWERF) in central Randolph County, WV (Fig. 1). The 3413-ha site is located
in the Unglaciated Allegheny Mountain and Plateau physiographic province
(Fenneman 1938) and ranges in elevation from 734 m to 1180 m. Average annual
precipitation ranged between 170 cm and 190 cm with an average snowfall >300
cm/year (National Oceanic and Atmospheric Administration 1998–2002). The
majority of the site was comprised of second-growth northern hardwood–Allegheny
hardwood forests (Keyser and Ford 2005). Forests were dominated by Fagus
grandifolia Ehrhart (American Beech), Acer rubrum L. (Red Maple), A. saccharum
Marshall (Sugar Maple), Liriodendron tulipfera L. (Yellow Poplar), and Prunus
serotina Ehrhart (Black Cherry). Other common species included Betula allegheniensis
Britton (Yellow Birch), Tilia americana L. (American Basswood), B. lenta
Figure 1. MeadWestvaco Wildlife and Ecosystem Research Forest (MWERF), Randolph
County, WV. Solid white polygons represent regenerating clearcuts (0–15 years since harvest)
during study period, 2006–2008.
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L. (Black Birch), and Quercus rubra L. (Northern Red Oak). Higher-elevation
areas were dominated by Picea rubens Sargent (Red Spruce) and Tsuga canadensis
Carriere (Eastern Hemlock) communities. Throughout much of the area, the understory
was composed of Smilax spp. (greenbrier) and Kalmia latifolia L. (Mountain
Laurel), with dense Rhododendron maximum L. (Rosebay Rhododendron) prevalent
along riparian areas. Dennstaedtia punctilobula Moore (Hay-scented Fern) also
was abundant throughout the understory due to excessive herbivory from historically
high Deer densities (Keyser and Ford 2005). Since 2000, more than 500 ha of
the MWERF have been harvested (Campbell et al. 2006).
Campbell et al. (2004) investigated the spatial ecology of Deer in the MWERF
during 1999–2002. At the time of that study, approximately 5% of the study area
was comprised of regenerating clearcuts (forest stands harvested within 15 years),
and the density of the Deer population was 12–20/km2 (Langdon 2001). During our
study (2006–2008), approximately 14% of the study area was comprised of regenerating
clearcuts and the density of the Deer population was 1.2–2.6/km2 (Crimmins
et al. 2013). Thus, the Deer population in the MWERF experienced an increase in
resource availability both through a reduction in the size of the population and an
increase in the absolute amount of forage (Miller et al. 2009). The causes for the
observed population decline were a combination of regional population declines,
likely resulting from historic overabundance and increasing predator populations,
and localized management actions that occurred between the conclusion of the
study by Campbell et al. (2004) and the onset of our study (Crimmins et al. 2012,
Miller et al. 2010).
Deer capture and monitoring
We captured Deer from January through March of 2005–2007 using modified
Clover traps (Clover 1954) baited with whole-kernel corn. We immobilized Deer
with an intramuscular injection of xylazine HCl (Sedazine, Fort Dodge Animal
Health, Fort Dodge, IA) at 2.2 mg/kg estimated body weight. We classified Deer
age as yearling (≤1.5 years) or adult (>1.5 years) according to tooth eruption and
wear patterns (Severinghaus 1949). We fit female Deer with VHF radio-collars
equipped with an 8-h mortality switch (Advanced Telemetry Systems, Isanti, MN).
We placed a uniquely numbered plastic ear tag on each animal for visual identification
(National Band and Tag, Newport, KY). We reversed sedation with a 50%
intramuscular and 50% subcutaneous injection of yohimbine HCl (Yobine, Lloyd
Laboratories, Shenandoah, IA) at 0.3 mg/kg (Wallingford et al. 1996). All capture
and handling methods were in accordance with the Animal Care and Use Committee
of West Virginia University (IACUC# 05-0706).
We attempted to locate Deer once per day, 3–4 d/week using biangulation (Mech
1983) or triangulation (White and Garrott 1990) from May 2006 to April 2008. We
used 3-element hand-held Yagi antennas and a TRX-1000 receiver (Wildlife Materials,
Murphysboro, IL) to locate collared animals. We recorded azimuths from ≥2
geo-referenced stations (n = 499) located throughout the MWERF with a maximum
of 15 min between azimuths to reduce the effect of animal movement on location
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accuracy (Schmutz and White 1990). Our telemetry methods were identical to those
used by Campbell et al. (2004). We assessed the accuracy of our telemetry locations
by placing 10 collars at geo-referenced locations throughout the study area during
July 2006 and collecting bearings from ≥10 of our geo-referenced stations to each
collar. We augmented telemetry locations with visual locations of radio-collared
animals identified via numbered ear tags recorded throughout the year. Preliminary
analyses indicated that our results were insensitive to the time of day when we
gathered locations (Barber-Meyer and Mech 2014).
Statistical analyses
We generated seasonal home-range and core-area estimates using the fixedkernel
method (Seaman and Powell 1996) with a least-squares cross-validated
bandwidth and reference grid-cell size (Gitzen et al. 2006, Kernohan et al. 2001).
We used 95% and 50% volume contours to define home ranges and core areas,
respectively. We calculated seasonal home ranges and core areas for animals
with ≥30 locations in a season (Seaman et al. 1999). We defined seasons as summer
(May–September), fall (October–December), and winter (January–April).
All of our home-range estimation procedures followed Campbell et al. (2004).
We detected no differences in home-range size between years (2006–2007 vs.
2007–2008) and subsequently pooled data between years. We tested for seasonal
differences in mean home-range size and core-area size using an analysis of variance
model with season as a treatment effect. We compared variability in homerange
and core-area sizes between seasons using 2-sided F-tests with Bonferroni
corrections. We estimated seasonal philopatry following Lesage et al. (2000) by
calculating the overlap in home-range and core-area polygons for individual Deer
in successive years. Using t-tests, we compared our seasonal home-range and
core-area size estimates and philopatry measurements to those previously found
for high density Deer populations (Campbell et al. 2004). We pooled our data
across age classes for all analyses. Because we pooled data across age classes, the
values we present for those observed at high population-density differ from those
presented in Campbell et al. (2004), wherein results were presented separately for
each age class.
Results
From May 2006 to April 2008, we collected 5252 locations from 35 individuals.
Of those, 33 had a sufficient number of locations (≥30) to generate at least
one seasonal home-range, resulting in 111 seasonal home ranges using a total of
4768 locations. Telemetry error in our trials of geo-referenced collars was minimal
(mean = 1.1°, n = 120). The number of seasonal home-ranges calculated per
individual ranged from 1 to 6. We were able to estimate seasonal philopatry from
15, 10, and 12 individuals in summer, fall, and winter, respectively. Our analysis
of variance indicated that mean home-range size was similar among seasons (F2 =
2.84, P = 0.06), as was mean core-area size (F2 = 1.73, P = 0.18) (Table 1). HomeNortheastern
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range size was less variable among individuals during winter than summer (F40,34 =
3.20, P < 0.01) or fall (F32,34 = 3.31, P < 0.01) (Table 1). Variability in home-range
size was similar in summer and fall (F32,40 = 0.966, P = 0.93) (Table 1). Similarly,
core-ranges were less variable in winter than summer (F40,34 = 3.60, P < 0.01) or
fall (F32,34 = 3.44, P < 0.01) (Table 1). As with home ranges, variability in core-area
size was similar in summer and fall ( F32,34 = 1.05, P = 0.88; Table 1).
Home-range estimates were larger after the reduction in Deer density and increase
in timber harvest than prior to reduction in density and increase in timber
harvest during summer (t43.68 = 6.55, P < 0.001), fall (t35.76 = 5.60, P < 0.001), and
winter (t71.51 = 3.69, P < 0.001) (Table 1). Similarly, core-area estimates were also
significantly larger in summer (t43.68 = 6.236, P < 0.001), fall (t35.76 = 5.475, P less than
0.0001), and winter (t71.51 = 5.463, P < 0.0001) than prior to population reduction
(Table 1). In general, seasonal home-range and core-area estimates were 2–4 times
greater than previously observed for Deer populations with high densities (Table 1;
Campbell et al. 2004).
Home-range philopatry did not differ among seasons (F = 1.57, df = 2; P = 0.22;
Table 2). Home-range philopatry was higher at low population-density than at high
population-density in summer (t = 4.084, df = 64.48; P < 0.0001), fall (t = 10.134,
df = 47.47; P < 0.0001), and winter (t = 9.277, df = 80.87, P < 0.0001) (Table 2).
Core-area philopatry differed among all seasons (F = 50.63, df = 2; P < 0.001),
with the highest values in fall and lowest in winter (Table 2). Core-area philopatry
at low density was similar to that at high density in summer (t = -1.20, df = 70.24;
P = 0.23), but was greater in fall (t = -13.31, df = 19.61, P < 0.001) and winter (t =
-2.58, df = 28.43; P = 0.02) (Table 2).
Table 1. Mean seasonal core-area (CA) and home-range (HR) sizes in hectares (95% CI) of female
White-tailed Deer on the MeadWestvaco Wildlife and Ecosystem Research Forest during periods of
high (1999–2002) and low (2006–2008) population density. Data from 1999–2002 was recalculated
from Campbell et al. (2004).
1999–2002 2006–2008
Season n Core area Home range n Core area Home range
Summer 165 18.0 (15.4–20.6) 99.5 (84.1–114.9) 42 87.4 (65.7–109.0) 389.7 (304.2–475.2)
Fall 162 18.6 (13.7–23.5) 104.9 (82.2–127.7) 33 89.7 (64.7–114.6) 383.4 (288.6–478.3)
Winter 177 26.1 (20.5–31.7) 152.1 (117.7–186.5) 36 64.6 (51.9–77.2) 267.6 (216.9–318.4)
Table 2. Mean seasonal core-area (CA) and home-range (HR) philopatry (95% CI), as measured in
percent overlap between years, of female White-tailed Deer on the MeadWestvaco Wildlife and Ecosystem
Research Forest during periods of high (1999–2002) and low (2006–2008) population density.
Data from 1999–2002 was recalculated from Campbell et al. (2004).
1999–2002 2006–2008
Season n Core area Home range n Core area Home range
Summer 59 40.2 (39.7–40.8) 55.8 (55.3–56.3) 15 44.8 (43.8–45.8) 66.1 (65.1–67.1)
Fall 58 18.2 (17.6–18.8) 41.2 (40.7–41.7) 10 74.0 (72.8–75.1) 69.2 (68.1–70.4)
Winter 83 26.2 (25.8–26.7) 51.2 (50.8–51.6) 12 35.7 (34.6–36.8) 68.8 (67.7–69.8)
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Discussion
It is widely recognized that population density can have substantial effects on
White-tailed Deer biology (Keyser et al. 2005a, 2005b; Kilpatri ck et al. 2001). Understanding
how density affects home-range patterns in Deer can aid managers in
developing management strategies to minimize disease transmission and herbivory
impacts to biodiversity and/or forest resources. Contrary to previous work suggesting
that seasonal changes in resource availability can affect space-use patterns in
White-tailed Deer (Labisky and Fritzen 1998), we found no evidence of difference
in home-range or core-area size among seasons in this study. Although previous research
has found similar consistencies among seasonal home-range patterns (e.g.,
Sargent and Labisky 1995), this result emphasizes that the effects of density on space
use can be difficult to predict, particularly when coupled with other factors such as
habitat quality and resource abundance. At high population-densities, Deer in the
MWERF exhibited seasonal variation in space-use, with larger home ranges and core
areas in winter than in summer or fall (Campbell et al. 2004). Changes in resource
availability in the MWERF led to species-specific changes in browsing rates (Crimmins
et al. 2010, Miller et al. 2009). Thus, seasonal variability in home-range sizes
previously observed (Campbell et al. 2004) may have been the result of seasonal-resource
limitation at high population densities and lower resource abundance, which
would be unlikely to have affected the population during our study due to reduced
population density and increased forage abundance. Additionally, we found higher
levels of site fidelity at low population densities, suggesting that per capita resource
limitation at high densities may have caused Deer to exhibit annual switches in
home-range areas in search of adequate resources. However, the increased site fidelity
that we observed could be an artifact of increased home-range sizes.
Timber harvesting on our study site led to a nearly 3-fold increase in the area
of early successional habitat. An increase in regenerating clearcuts from approximately
5% of the study area in 1999–2002 to nearly 14% in 2006–2008 increased
the amount of browse available to deer during spring and summer, a period of high
nutritional requirements for lactating females (Ford et al. 1993, Wentworth et al.
1990). Regenerating clearcuts at our study area contained abundant forage compared
to mature forest stands (Crimmins et al. 2010), highlighting the ecological
importance of this component of the landscape. Home-range and core-area sizes
were approximately 300% larger than prior to the population reduction likely due
to a combination of reduced population density and increased habitat disturbance.
Research in other regions has indicated that population density and habitat patchiness
can affect home-range size independently (Ford 1983, Kilpatrick et al. 2001,
Lopez et al. 2005). Our finding indicating that Deer had larger home ranges at decreased
population densities is contrary to several previous studies (Kilpatrick et
al. 2001, McNulty et al. 1997) and supports the theory that decreased intraspecific
competition is a driving factor in structuring home-range patterns.
Our finding of increased site fidelity at lowered densities contrasts with previous
research focused on socio-spatial patterns in Deer family groups (McNulty et al.
1997). It is possible that these increases in site fidelity were the result of increases
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2015 Vol. 22, No. 2
in per capita resources, whereby abundant resources reduced the need for exploratory
behavior in search for adequate forage. This explanation is supported by previous
research at our study area that documented an increase in the abundance of forage
following the reduction in Deer population density and increase in timber harvest
(Crimmins et al. 2010). Although increases in the availability of resources could
lead to the changes in home-range patterns we observed, alternative explanations
such as increased predator populations also merit consideration. For example, the
substantial increases in Canis latrans Say (Coyote) populations that have occurred
in the region (Crimmins et al. 2012) could lead to altered space-use patterns by Deer
in an attempt to reduce predation risk. Because of the coupled changes in population
density and habitat structure that occurred during our study, it is impossible
to determine the absolute influence of either factor on the changes in home-range
patterns we observed. Additional research to examine changes in habitat-use patterns
could further elucidate the mechanisms behind these observed changes in
home-range patterns. Previous research has shown that overall browsing rates
in the MWERF declined following the Deer population decline and increase in
timber harvest, but that browsing rates for forage species generally thought to be
preferred by White-tailed Deer showed a negligible decline (Crimmins et al. 2010).
This result suggests that increases in home-range size may have been a function of
Deer actively searching for preferred food sources over greater areas rather than
relying on sub-optimal forage resources within a smaller area. Regardless of the
causal mechanism behind the observed changes in home-range patterns, our results
suggest that in this region, it may be difficult to successfully implement management
strategies designed to reduce White-tailed Deer abundance at a localized scale
(Campbell et al. 2004).
Acknowledgments
Funding and support for this project was provided by the MeadWestvaco Corporation,
Penn-Virginia Resources, the West Virginia Division of Natural Resources, and the Division
of Forestry and Natural Resources at West Virginia University.
Literature Cited
Barber-Meyer, S.M., and L.D. Mech. 2014. Accuracy of estimating White-tailed Deer
diel May–June home ranges using only daytime locations. Wildlife Biology in Practice
10:62–68.
Brown, J.L. 1969. Territorial behavior and population regulation in birds. Wilson Bulletin
81:293–329.
Burt, W.H. 1943. Territoriality and home-range concepts as applied to mammals. Journal of
Mammalogy 24:346–352.
Campbell, T.A., B.R. Laseter, W.M. Ford, and K.V. Miller. 2004. Feasibility of localized
management to control White-tailed Deer in forest-regeneration areas. Wildlife Society
Bulletin 32:1124–1131.
Campbell, T.A., B.R. Laseter, W.M. Ford, R.H. Odom, and K.V. Miller. 2006. Abiotic factors
influencing deer browsing in West Virginia. Northern Journal of Applied Forestry
23:20–26.
Northeastern Naturalist Vol. 22, No. 2
S.M. Crimmins, J.W. Edwards, T.A. Campbell, W.M. Ford, P.D. Keyser, B.F. Miller, and K.V. Miller
2015
411
Clover, M.R. 1954. A portable deer trap and catch-net. California Fish and Game
40:367–373.
Clutton-Brock, T.H. 1989. Mammalian mating systems. Proceedings of the Royal Society
of London B-Biological Sciences 236:339–372.
Crimmins, S.M., J.W. Edwards, W.M. Ford, P.D. Keyser, and J.M. Crum. 2010. Browsing
patterns of White-tailed Deer following increased timber harvest and a decline in population
density. International Journal of Forestry Research 2010:ID592034
Crimmins, S.M., J.W. Edwards, and J.M. Houben. 2012. Canis latrans (Coyote) habitat use
and feeding habits in central West Virginia. Northeastern Naturalist 19:411–420.
Crimmins, S.M., J.W. Edwards, P.D. Keyser, J.M. Crum, W.M. Ford, B.F. Miller, T.A.
Campbell, and K.V. Miller. 2013. Survival rates of female White-tailed Deer on an
industrial forest following a decline in population density. Proceedings of the Central
Hardwood Forest Conference. 117:487–496.
Fenneman, N.M. 1938. Physiography of the Eastern United States. McGraw-Hill, New
York, NY. 714 pp.
Ford, R.G. 1983. Home range in a patchy environment: Optimal-foraging predictions.
American Zoologist 23:315–326.
Ford, W.M., A.S. Johnson, P.E. Hale, and J.W. Wentworth. 1993. Availability and use of
spring and summer woody browse by deer in clearcut and uncut forests of the southern
Appalachians. Southern Journal of Applied Forestry 17:116–119.
Fretwell, S.D., and J.S. Calver. 1969. On territorial behavior and other factors influencing
habitat distribution in birds. Acta Biotheriologica 19:37–44.
Garroway, C.J., and H.G. Broders. 2005. The quantitative effects of population density and
winter weather on the body condition of White-tailed Deer (Odocoileus virginianus) in
Nova Scotia, Canada. Canadian Journal of Zoology 83:1246–1256.
Getz, L.L. 1961. Home ranges, territoriality, and movement of the Meadow Vole. Journal
of Mammalogy 42:24–36.
Gitzen, R.A., J.J. Millspaugh, and B.J. Kernohan. 2006. Bandwidth selection for fixedkernel
analysis of animal-utilization distributions. Journal of Wildlife Management
70:1334–1344.
Henderson, D.W., R.J. Warren, J.A. Cromwell, and R.J. Hamilton. 2000. Responses of urban
deer to a 50% reduction in local herd density. Wildlife Society Bulletin 28:902–910.
Kernohan, B.J., R.A. Gitzen, and J.J. Millspaugh. 2001. Analysis of animal-space use and
movements. Pp.125–166, In J.J. Millspaugh and J. Marzluff (Eds.). Radio Tracking and
Animal Populations. Academic Press, San Diego, CA. 474 pp.
Keyser, P.D., and W.M. Ford. 2005. Ten years of research on the MeadWestvaco Wildlife
and Ecosystem Research Forest: An Annotated Bibliography. USDA Forest Service.
General Technical Report, GTR NE-330. Northeastern Research Station, Newtown, PA.
Keyser, P.D., D.C. Guynn, and H.S. Hill, Jr. 2005a. Density-dependent recruitment patterns
in White-tailed Deer. Wildlife Society Bulletin 33:222–232.
Keyser, P.D., D.C. Guynn, and H.S. Hill, Jr. 2005b. Population density–physical condition
relationships in White-tailed Deer. Journal of Wildlife Management 69:356–365.
Kilpatrick, H.J., S.M. Spohr, and K.K. Lima. 2001. Effects of population reduction on
home ranges of female White-tailed Deer at high densities. Canadian Journal of Zoology
79:949–954.
Labisky, R.F., and D.E. Fritzen. 1998. Spatial mobility of breeding female White-tailed
Deer in a low-density population. Journal of Wildlife Management 62:1329–1334.
Langdon, C.A. 2001. A comparison of White-tailed Deer population estimation methods in
West Virginia. M.Sc. Thesis. West Virginia University, Morgantown, WV. 130 pp.
Northeastern Naturalist
412
S.M. Crimmins, J.W. Edwards, T.A. Campbell, W.M. Ford, P.D. Keyser, B.F. Miller, and K.V. Miller
2015 Vol. 22, No. 2
Lesage, L., M. Crete, J. Hout, A. Dumont, and J. Ouellet. 2000. Seasonal home-range size
and philopatry in two northern White-tailed Deer populations. Canadian Journal of Zoology
78:1930–1940.
Lopez, R.R., P.M. Harveson, M.N. Peterson, N.J. Silvy, and P.A. Frank. 2005. Changes in
ranges of Florida Key Deer: Does population density matter? Wildlife Society Bulletin
33:343–348.
McNulty, S.A., W.F. Porter, N.E. Matthews, and J.A. Hill. 1997. Localized management for
reducing White-tailed Deer populations. Wildlife Society Bulletin 25:264–271.
Mech, L.D. 1983. Handbook of Animal Radio-Tracking. University of Minnesota Press,
Minneapolis, MN. 120 pp.
Miller, B.F., T.A. Campbell, B.R. Laseter, W.M. Ford, and K.V. Miller. 2009. White-tailed
Deer herbivory and timber-harvesting rates: Implications for regeneration success. Forest
Ecology and Management 258:1067–1072.
Miller, B.F., T.A. Campbell, B.R. Laseter, W.M. Ford, and K.V. Miller. 2010. A test of
localized management for reducing White-tailed Deer herbivory in central Appalachian
regeneration sites. Journal of Wildlife Management 74:370–378.
National Oceanic and Atmospheric Administration. 1998–2002. Climatological data, West
Virginia. Volumes 106–110. Environmental Data Service, Ashville, NC.
Sanderson, G.C. 1966. The study of mammal movements: A review. Journal of Wildlife
Management 30:215–235.
Sargent, R.A., and R.F. Labisky. 1995. Home range of male White-tailed Deer in hunted
and non-hunted populations. Proceedings of the Southeastern Association of Fish and
Wildlife Agencies 49:389–398.
Schmutz, J.A., and G.C. White. 1990. Error in telemetry studies: Effects of animal movement
on triangulation. Journal of Wildlife Management 54:506–510.
Seaman, D.E., and R.A. Powell. 1996. An evaluation of the accuracy of kernel-density estimators
for home-range analysis. Ecology 77:2075–2085.
Seaman, D.E., J.J. Millspaugh, B.J. Kernohan, G.C. Brundige, K.J. Raedeke, and R.A. Gitzen.
1999. Effects of sample size on kernel home-range estimates. Journal of Wildlife
Management 63:739–747.
Severinghaus, C.A. 1949. Tooth development and wear as criteria of age in White-tailed
Deer. Journal of Wildlife Management 13:195–216.
Tiersen, W.C., G.F. Mattfield, R.W. Sage, Jr., and D.F. Behrend. 1985. Seasonal movements
and home ranges of White-tailed Deer in the Adirondacks. Journal of Wildlife Management
49:760–769.
VanHorne, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife
Management 49:92–103.
Wallingford, B.D., R.A. Lancia, and E.C. Soutiere. 1996. Antagonism of xylazine in Whitetailed
Deer with intramuscular injection of yohimbine. Journal of Wildlife Diseases
32:399–402.
Wentworth, J.M., A.S. Johnson, P.E. Hale, and K.E. Kammermeyer. 1990. Seasonal use of
clearcuts and food plots by White-tailed Deer in the southern Appalachians. Proceedings
of the Southeastern Association of Fish and Wildlife Agencies 44:215–223.
White, G.C., and R.A. Garrott. 1990. Analysis of Wildlife Radio-Tracking Data. Academic
Press, San Diego, CA. 383 pp.
Williams, S.C., and A.J. DeNicola. 2002. Home-range increase of lactating female Whitetailed
Deer following herd reduction. Northeast Wildlife 57:29–38.