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Canis latrans (Coyote) Habitat Use and Feeding Habits in Central West Virginia
Shawn M. Crimmins, John W. Edwards, and John M. Houben

Northeastern Naturalist, Volume 19, Issue 3 (2012): 411–420

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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. 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. 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