2012 NORTHEASTERN NATURALIST 19(3):411–420
Canis latrans (Coyote) Habitat Use and Feeding Habits in
Central West Virginia
Shawn M. Crimmins1,2, John W. Edwards1, and John M. Houben3
Abstract - Canis latrans (Coyote) populations are expanding throughout the
eastern United States, making them the apex predator in many systems. Despite
abundant research in the western United States, relatively little information exists
on the space use or feeding patterns of Coyotes in the forested landscapes of the
Appalachians. We used radio-telemetry and scat analysis to describe seasonal habitat
use and feeding patterns of Coyotes in central West Virginia during 2006–2008.
Odocoileus virginianus (White-tailed Deer) was the most common prey, occurring
in 76% of scats collected in winter and 45% of scats collected in summer. Rodents
were the most common prey item in summer, occurring in 48% of scats; other prey
items occurred in <20% of scats. Coyotes selected for recently harvested forest
stands while avoiding intact stands in both summer and winter. Despite exhibiting
seasonal prey-switching behavior, Coyotes in this region do not alter habitat-use
patterns with respect to season. Coyotes in our study seem to be opportunistic feeders
that prefer areas with abundant cover. Their opportunistic feeding patterns may
contribute to their rapid population expansion in this region.
Introduction
Canis latrans Say (Coyote) populations are expanding rapidly in the eastern
United States (Lovell et al. 1998), making them a top predator in many areas
(Gompper 2002). However, little is known of the ecology of Coyotes in the eastern
United States compared to populations in the western United States. The expansion
of eastern Coyote populations has been largely facilitated by colonization from
northern and western Coyote populations (Bozarth et al. 2011) and by hybridization
with western Coyotes and Canis lupus lycaon L. (Eastern Wolves) (Kays et al.
2010, Way et al. 2010), making these populations somewhat unique compared to
more intensively studied populations in the western United States. Despite recent
increases in Coyote population size throughout the region, it has been suggested
that northeastern forests provide marginal habitat for Coyotes (Crete et al. 2001).
Thus, there seems to be a disconnect between predictions of Coyote population
dynamics in this region and observed changes in population size. Recent investigations
of Coyote populations in suburban (Gehrt et al. 2009, Morey et al. 2007)
and agricultural (Kamler and Gipson 2000) landscapes have provided insights into
1Division of Forestry and Natural Resources, West Virginia University, Morgantown,
WV 26506. 2Current address - 413A Charles Clapp Building, University of
Montana, Missoula, MT 59802. 3USDA Animal and Plant Health Inspection Service,
Wildlife Services, Cottageville, WV 25239. *Corresponding author - shawn.
crimmins@umontana.edu.
412 Northeastern Naturalist Vol. 19, No. 3
their ecology, but have yielded little consensus on patterns of space-use or feeding
habits of Coyotes. In heavily forested landscapes, Coyotes may exhibit substantially
different patterns of space use and feeding habits (Patterson and Messier 2001)
than observed in suburban or agricultural landscapes.
Coyotes exhibit large variation in home-range size and habitat-use patterns.
Reported Coyote home ranges vary in size from 4.1 km2 (Kamler and Gipson 2000)
in Kansas to 68.7 km2 (Litvaitis and Shaw 1980) in Oklahoma, despite similar
habitats. Home-range size in Coyotes can also vary by age, gender, and season
(Holzman et al. 1992), making generalities diffi cult. Similarly, habitat use by Coyotes
can exhibit substantial variability. Kamler and Gipson (2000) reported that
resident Coyotes selected open grassland habitats more than expected in Kansas.
Conversely, Chamberlain et al. (2000) found that Coyotes avoided open habitats
in Mississippi, highlighting the lack of congruence in space-use patterns among
populations. However, little information exists on the use of differing forestcover
types by Coyotes inhabiting forested landscapes of the central Appalachians.
Coyote space-use patterns can also vary as a function of prey availability, with
home-range size decreasing as prey abundance increases (Mills and Knowlton
1991, Patterson and Messier 2001). Recently, however, Boser (2009) found that
Coyote movements were unrelated to prey densities in forested and agricultural
landscapes of New York. Most dietary studies suggest that Coyotes are opportunistic
feeders, increasing their use of specifi c prey items as they become more
prevalent (Bartel and Knowlton 2005, Harrison and Harrison 1984, Litvaitis and
Shaw 1980). However, several studies have found that Coyote feeding patterns do
not follow patterns of prey abundance, indicating a preferential use of primary prey
items (Morey et al. 2007, Patterson et al. 1998). For example, Boser (2009) found
that Odocoileus virginianus Zimmerman (White-tailed Deer) was the dominant
food among Coyotes in forested areas of New York. Previous studies have indicated
Coyotes readily feed on White-tailed Deer and rodents, regardless of response
to variation in prey abundance (Chamberlain and Leopold 1999, Hidalgo-Mihart
et al. 2001). Coyotes also exhibit seasonal variability in food habits (Andelt et al.
1987), often resulting from variable prey densities or winter weather patterns. The
objectives of our study were to describe seasonal space use and feeding patterns of
Coyotes in a heavily forested landscape of northeastern North America where Coyote
populations are thought to be expanding.
Methods
Study area
We conducted our study on the MeadWestvaco Wildlife and Ecosystem Research
Forest (MWERF) in central Randolph County, WV from May 2006 to
April 2008. The 3413-ha site ranges in elevation from 734 to 1180 m. Average
annual precipitation on the site ranges between 170 and 190 cm, with an average
snowfall >300 cm/year. The majority of the site was logged between 1916
and 1928 and is now comprised primarily of second-growth northern hardwood-
Allegheny hardwood forests (Keyser and Ford 2005). The forest communities
of the MWERF are dominated by Fagus grandifolia Ehrhart (American Beech),
2012 S.M. Crimmins, J.W. Edwards, and J.M. Houben 413
Prunus serotina Ehrhart (Black Cherry), various Acer spp. (maple), Betula allegheniensis
Britt. (Yellow Birch), and Quercus rubra L. (Northern Red Oak).
High-elevation areas were dominated by Picea rubens Sargent (Red Spruce)
and Tsuga canadensis Carriere (Eastern Hemlock) communities. At lower elevations,
Tilia americana L. (American Basswood), B. lenta L. (Black Birch), and
Liriodendron tulipifera L. (Yellow Poplar) are also present. Throughout much of
the area, the understory is dominated by Smilax rotundifolia L. (Greenbriar) and
Kalmia latifolia L. (Mountain Laurel), with Rhododendron maximum L. (Rosebay
Rhododendron) prevalent along riparian areas. Dennstaedtia punctilobula
Moore (Hay-scented Fern) is also abundant throughout the understory due to
excessive herbivory from historically high White-tailed Deer densities (Keyser
and Ford 2005). Since 2000, more than 500 ha of forest have been harvested on
the MWERF, of which 75% has been clearcut, with the remaining 25% in deferment
cuts, diameter-limit cuts, and marked-selection cuts. Harvest units have
averaged 15 ha in size since the mid-1990s. Coyote populations in West Virginia
are generally thought to be rapidly expanding. Annual surveys of hunters in West
Virginia conducted by the West Virginia Division of Natural Resources indicate
that Coyote sightings across the state have increased substantially between 1995
and 2005 (fig. 1).
Coyote capture and telemetry
We captured Coyotes in May and October 2006 using Number 3 Victor softcatch
foothold traps (Woodstream, PA) baited with commercial and homemade
lure and Coyote urine. We placed traps along roads at locations with recent Coyote
sign (track or scats) or where other items (e.g., gut piles from harvested deer)
would attract Coyotes. Upon capture, we physically restrained Coyotes with a
figure 1. Number
of Coyotes seen per
1000 hours of hunting
by fall archery
hunters and spring
turkey hunters in
West Virginia from
1995–2005. Dotted
line represents signifi
cant (P < 0.001,
β = 0.581) linear
trend.
414 Northeastern Naturalist Vol. 19, No. 3
catch pole and chemically immobilized them with an intramuscular injection
of ketamine and xylazine (6.6 mg/kg ketamine + 2.2 mg/kg xylazine; Beheler-
Amass et al. 1998). We recorded the sex, weight, body length, and approximate
age (juvenile/adult) of each Coyote. We placed a mortality-sensitive radio collar
(Advanced Telemetry Systems, Ishanti, MN) and numbered plastic ear tags (National
Band and Tag, Newport, KY) on each Coyote. We administered parvovirus
and canine distemper vaccinations prior to release. We released all Coyote at
their capture locations.
We located radio-collared Coyotes using biangulation and triangulation
(White and Garrott 1990) from geo-referenced (n = 499) stations located
throughout the study area. We attempted to locate Coyotes once per day, 1–2
days per week from May 2006 to April 2008. Location estimates were generated
using program LOCATE II (Nams 2006). To reduce location error, we only used
azimuths between 45 and 135 degrees (Springer 1979) and taken within 15 minutes
of each other (Schmutz and White 1990). Animals were tracked until death,
study termination, or loss of radio contact.
Habitat use
We classifi ed forest stands in our study area as clearcut (timber harvest within
10 yr.) or forested (no timber removal within 10 yr.) cover types. We compared the
relative use, measured by number of telemetry locations of each individual within
a forest-cover type, and availability of each forest-cover type across the study area
using selection ratios with a Type II study design (Manly et al. 2002) within a GIS.
We chose not to analyze habitat selection within individual home ranges (2nd order
selection; Johnson 1980) because several animals lacked a suffi cient number of telemetry
locations to generate home-range estimates (Seaman et al. 1999). Selection
ratios > 1 indicate selection for a resource whereas ratios < 1 indicate avoidance.
We only included animals for which we had ≥10 locations throughout the season.
Preliminary analyses indicated that space-use patterns did not qualitatively differ
among years, gender, or age class, although sample sizes were too small to make
robust statistical comparisons. Therefore, we pooled data across these factors and
only compared selection ratios between seasons. All analyses were conducted in the
R programming language (R Development Core Team 2010).
Scat collection and analysis
We opportunistically collected scats deposited along a predetermined 20-km
network of roads in our study area throughout the year. Although varying sampling
effort among seasons and years led to varying sample size, our collection
effort was suffi cient for comparisons between seasons. We defi ned seasons as
summer (May–Sep) and winter (Oct–Apr) (Kamler et al. 2005). Prior to the start
of each season, we removed all remaining scats from the entire road network to
ensure that all scats were deposited during our target season. Once we located a
scat, it was placed in a plastic bag and frozen for storage and transport. We dried
scats at approximately 60 °C for ≥24 h prior to analysis (Kelly and Garton 1997).
We used visual identifi cation of prey remains and structural characteristics of
hairs (Moore et al. 1997) to defi ne 7 categories of prey (deer, rodent, lagomorph,
2012 S.M. Crimmins, J.W. Edwards, and J.M. Houben 415
avian, herpetofauna [amphibian or reptile], unknown mammal, plant). We expressed
Coyote use of prey using frequency of occurrence (Litvaitis et al. 1994).
We compared the proportion of scats containing each prey category between
seasons using a chi-square test. We compared proportions among prey categories
within each season using pairwise chi-square tests (Zar 1998). Chi-square tests
indicated similar proportions of prey items between years, therefore we pooled
data across years for all seasonal comparisons. Statistical signifi cance was accepted
at α = 0.05.
Results
Habitat use
We captured and radio-collared 7 Coyotes (4F: 3M) during our study. From
these, we were able to gather suffi cient telemetry data to estimate 9 seasonal selection
ratios from 6 individuals over the two years of our study (4F: 2M, x̅ = 24.9
± 3.4 locations/season). Selection ratios indicated that Coyotes exhibited strong
figure 2. Seasonal
selection ratios for
clearcut (A) and
forested (B) cover
types by Coyotes
in central West Virginia.
Black bars
represent 95% confi
dence interval.
416 Northeastern Naturalist Vol. 19, No. 3
selection for clearcuts (fig. 2A) and strong avoidance of forested areas (fig. 2B)
in both summer and winter. There were no seasonal differences in selection ratios
for either cover type based on confi dence interval overlap.
Feeding habits
We collected 128 scats (n = 86 summer, n = 42 winter), with 83 collected in the
fi rst year and 45 collected in the second year of our study. Deer remains occurred
more often (76%) than any other prey item during winter (P < 0.001), with all other
prey items occurring in less than 25% of scats in winter (fig. 3). Deer and rodent
remains did not differ in frequency of occurrence during summer (P = 0.879), but
both occurred more frequently than any other prey item (P < 0.001). Deer remains
were signifi cantly more common in winter than summer (P = 0.001), whereas rodent
remains were signifi cantly more common in summer than winter (P = 0.012;
fig. 3). No other prey items differed in frequency of occurrence between seasons,
with all occurring in less than 20% of scats in each season (fig. 3).
Discussion
Coyotes in our study showed a strong selection for areas with recent timber
harvests. In our study site, these areas are characterized by abundant understory
plant growth relative to the surrounding mature forest and are readily used by
figure 3. Proportion of Coyote scats (n = 128) containing specifi c prey items in summer
(gray bar) and winter (black bar). Prey items marked with asterisk indicate proportions
differed signifi cantly (P < 0.05) between seasons.
2012 S.M. Crimmins, J.W. Edwards, and J.M. Houben 417
deer for foraging (Crimmins et al. 2010). Additionally, these areas have an
abundance of logging roads and trails from timber harvesting operations, which
can increase carnivore predation rates on ungulates (Merrill et al. 2010). The
areas with intact timber that were avoided by Coyotes were characterized by a
distinct lack of ground cover, which may have reduced the abundance of prey
items such as small mammals in these areas. Boser (2009) concluded that differences
in cover-type selection by Coyotes in New York were best explained
by mortality risk rather than foraging opportunities, whereas studies in other
regions indicate that Coyote habitat use is strongly related to prey availability
(e.g., Mills and Knowlton 1991). It is possible that clearcuts within our study
area contained greater densities of primary (deer) and secondary (rodents)
prey items than mature forest areas, suggesting that prey availability may be
an important driving factor in space-use patterns by Coyotes in this region, although
more detailed studies of Coyote movements and detailed data on prey
abundance are required to reach any conclusions and may be an appropriate objective
for future research.
Deer remains were found in 76% of winter scats and 45% of summer scats, indicating
a strong reliance on deer, particularly during the winter period. Coyotes
commonly feed on deer, and previous research has documented similar patterns
of deer remains in Coyote scats in northeastern North America (Patterson et al.
1998). We do not know if the increase in prevalence of deer remains during the
winter was the result of increased predation on deer or scavenging of carcasses,
as Coyotes will readily scavenge carrion (Boser 2009, Chamberlain and Leopold
1999). Because deer in our study area were subject to human harvest (Crimmins
et al. in press), Coyotes’ feeding patterns during the hunting season may have
reflected the availability of discarded gut piles or other hunting-related carrion.
Adult deer on our study area exhibited high survival rates during summer (Campbell
et al. 2005; Crimmins et al., in press), suggesting that deer remains found
in summer scats likely resulted from predation of deer fawns, which has been
observed elsewhere in the region (Vreeland et al. 2004). These two factors indicate
that Coyotes in our study area are not readily preying on adult deer, but are
instead relying on temporal shifts in the availability of carrion and fawns. These
results support fi ndings in other regions suggesting that Coyotes exhibit opportunistic
feeding behavior and regularly switch primary prey items depending on
availability (Bartel and Knowlton 2005, Patterson et al. 1998). The abundance
of rodent remains found in scats during the summer also supports the hypothesis
that Coyotes will opportunistically feed on prey items as they are available. This
seasonal change in secondary prey items also has been documented for Coyotes
in the western United States (Bartel and Knowlton 2005). Although vegetative
forage was abundant on our study area (Crimmins et al. 2010), we found
that plant material comprised a relatively minor part of Coyotes’ diet. Previous
analyses of stomach contents from dead Coyotes have suggested that plant material
can be a more common component of Coyote diets in West Virginia (Wykle
1999). This discrepancy between our results and previous studies suggests that
further investigations of Coyote dietary patterns in this region are needed.
418 Northeastern Naturalist Vol. 19, No. 3
Our study was conducted in a region thought to have rapidly expanding Coyote
populations. One ecological benefi t of the presence and expansion of large carnivore
populations within the region is the potential for Coyotes to limit White-tailed
Deer populations, which are considered ecologically overabundant in much of
northeastern North America (McShea et al. 1997). Other ecological consequences,
such as changes in small-mammal community structure and abundance of other
mesocarnivores (Henke and Bryant 1999), could also occur if Coyote populations
in the region continue to expand. Because Coyotes can exhibit substantial
geographic variability in dietary and space-use patterns, and because there is very
little basic ecological information regarding Coyotes in this region, it is diffi cult
to predict the future dynamics of these expanding populations. Our results suggest
that Coyotes are extremely adaptable predators that can readily switch their
prey base depending on the availability of resources and can use cover types often
thought to be poor habitat. However, because our study was small in scale, care
should be taken not to assume that our results are representative of Coyote populations
throughout the region. Additional research is required to fully understand the
dynamics of these populations and their potential ecological effects in northeastern
North America.
Acknowledgments
We thank the MeadWestvaco Corporation, the Northeast Wildlife Damage Management
Cooperative, the West Virginia Division of Natural Resource, USDA-APHIS Wildlife
Services, and West Virginia University for providing support for this project. We are
grateful to the many technicians who assisted with data collection, especially C. Brabham
and J. Cecil for laboratory analyses. R. Tucker (WVDNR) graciously provided hunter
survey data.
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