2008 SOUTHEASTERN NATURALIST 7(1):135–144 Seasonal Food Habits of the Coyote in the South Carolina Coastal Plain Joshua D. Schrecengost1,*, John C. Kilgo2, David Mallard3, H. Scott Ray4, and Karl V. Miller5 Abstract - Spatial and temporal plasticity in Canis latrans (coyote) diets require regional studies to understand the ecological role of this omnivorous canid. Because coyotes have recently become established in South Carolina, we investigated their food habits by collecting 415 coyote scats on the Savannah River Site in western South Carolina from May 2005–July 2006. Seasonally available soft mast was the most common food item in 12 of the 15 months we sampled. Odocoileus virginianus (white-tailed deer) was the most common food item during December (40%) and March (37%). During May–June, fruits of Prunus spp. and Rubus spp. were the most commonly occurring food items. Fawns were the most common mammalian food item during May and June of both years despite low deer density. Introduction Canis latrans Say (coyote) is historically associated with western North America. However, during the last 50 years, they have expanded their range into the southeastern US, aided by humans and anthropogenic changes in the landscape (Hill et al. 1987). Coyote food habits have been explored in detail throughout much of their historical range (Andelt et al. 1987, Bekoff 1977, Murie 1945). However, because coyotes exhibit temporal and spatial variability in food item use (Bekoff 1977, Cypher et al. 1994), the findings of these studies have limited value in the Southeast. Coyote food habits have been documented in several areas of the Southeast (Blanton and Hill 1989, Gipson 1974, Hall 1979, Hoerath 1990, Lee 1986, Michaelson 1975, Smith and Kennedy 1983, Thornton et al. 2004, Wagner 1993, Wilson 1967, Wooding 1984), but no published record exists for South Carolina or adjacent states. Although there is considerable overlap in the types of food items documented in these studies, the importance of particular food items in the coyote diet differs geographically among study areas and temporally among seasons, probably due to changes in food-item availability. Coyote food habits can also be affected by changes in land-use practices (Gipson and Sealander 1976) and successional changes in vegetation (Andelt et al. 1987). The effect of coyote predation on game-animal populations has been of interest for decades. In the Southeast, only two studies have focused on specific game animals. Wagner (1993) studied coyote diet during the Meleagris gal- 1Georgia Department of Natural Resources, Wildlife Resources Division, 1401 Dean Street, Suite I, Rome, GA 30161. 2USDA Forest Service Southern Research Station, PO Box 700, New Ellenton, SC 29809. 3Building 5884, First Division Road, Fort Benning, GA 31905. 4USDA Forest Service-Savannah River, PO Box 700 New Ellenton, SC 29809. 5D.B. Warnell School of Forestry and Natural Resources, University of Georgia, Athens, GA 30602. *Corresponding author - josh_schrecengost@dnr.state.ga.us. 136 Southeastern Naturalist Vol.7, No. 1 lopavo Linnaeus (Wild Turkey) reproductive season in Arkansas, Mississippi, Alabama, and Florida, and reported Wild Turkey occurred in less than 2% of scats collected. Blanton and Hill (1989) compared diets of coyotes in areas of high and low Odocoileus virginianus Zimmerman (white-tailed deer) densities during the fawning period in Alabama, Kentucky, Mississippi, and Tennessee. Occurrence of deer in coyote scats and stomachs ranged from 9% in low-density areas to 74% in high-density areas during the fawning period. This wide variation in occurrence is consistent with the findings of other southeastern studies. Michaelson (1975) and Wilson (1967) both reported annual deer occurrences less than 5%, whereas Wooding (1984) reported deer occurrences of 71% during August. Variability in food habits, even among similar habitats, limits the use of prior studies in determining site-specific ecological impact of coyotes. Detailed data describing coyote use of intensely managed populations such as white-tailed deer are of particular interest. Therefore, our objectives were to determine the breadth of coyote food habits and identify seasonal trends in the coyote diet on the Savannah River Site in South Carolina. Study Area Our research was conducted on the 80,000-ha Savannah River Site (SRS) in Aiken, Barnwell, and Allendale counties, SC. The SRS is a US Department of Energy National Environmental Research Park located in the Upper Coastal Plain physiographic province along the Savannah River (Imm and McLeod 2005). General public access to the SRS is heavily restricted. Coyotes were first reported on the SRS in 1986, and since that time, the population has expanded dramatically (Mayer et al. 2005), and coyotes are now observed frequently. Topography of the SRS is gently rolling to fl at, and elevation ranges from 20–130 m. Once used for agriculture, SRS is now predominately forested (97%). Pinus palustris Miller (longleaf pine) and Pinus taeda Linnaeus (loblolly pine) dominate the overstory canopy (68%). Other major vegetative types include swamps and bottomland hardwood (22%) and upland hardwood (7%) forests (Imm and McLeod 2005). Approximately 12% of the total forest stands are <10 years of age (Blake and Bonar 2005). The SRS is intersected by over 2600 km of roads, logging trails, and railroads (Blake et al. 2005). The white-tailed deer population on the SRS is managed to limit deer-vehicle collisions and deer impact on the environment. Controlled dog hunting is used to maintain a pre-hunt target population of 4000 deer (Johns and Kilgo 2005). Currently, deer population density is estimated at 1 per 26 to 29 ha (H.S. Ray and J.C. Kilgo, unpubl. data). Hunts during the course of this study occurred October through December 2005. In recent years, data collected from the hunts on SRS show a dramatic decline in harvest of fawns and yearlings despite no changes in hunt methodology (H.S. Ray, unpubl. data). This suggests that white-tailed deer recruitment is declining at SRS, coincident with increasing coyote abundance. The Sus scrofa Linnaeus (wild hog) population on SRS was estimated at 900 animals in 2003 (Mayer 2005). Wild hogs are hunted and trapped year round to minimize impact on the environment and vehicle collisions. From May 2005 2008 J.D. Schrecengost, J.C. Kilgo, D. Mallard, H.S. Ray, and K.V. Miller 137 through June 2006, 172 hogs were killed on SRS. Most wild hog carcasses are disposed of on site, which provides an incidental source of carrion for coyotes. Methods We collected coyote scats opportunistically along roads and rights-of-way throughout SRS from May 2005 through July 2006. Especially large or small scats were not collected to avoid inclusion of scats of Canis familiaris Linnaeus (domestic dog) and Urocyon cinereoargenteus Schreber (gray fox). Scats were subjectively evaluated in the field for relative moisture content and decomposition in an attempt to collect samples <5 days old. Scats were placed in plastic bags labeled with date and location of collection and stored frozen to minimize decomposition. For analysis, scats were oven dried at 65 °C for 72–96 hours, and food items were separated manually. We identified food items macroscopically from hair, tooth, claw, and hoof fragments, as well as plant residues. Dorsal guard hairs were identified microscopically when necessary using the pigment patterns of the medulla (Moore et al. 1974) and compared to reference slides prepared from the University of Georgia Warnell School of Forestry and Natural Resources mammalian collection. White-tailed deer fawn remains were separated from adult deer remains whenever possible by the physical characteristics of the hairs as well as the presence of small hooves. In cases of uncertain age class, the deer remains were included as adult. Plant residues, primarily seeds, were compared to reference manuals and collections of known specimens housed at the Warnell School of Forestry and Natural Resources. We recorded the occurrence as well as a visual estimate of percentage volume for each food item in each scat. Previous studies have used several terms interchangeably to describe the occurrence of food items in the diet of coyotes (Wagner 1993). We chose to define “percent of scats” as the percent of a sample of scats in which a food item occurs (Table 1), and “percent of occurrence” as the number of times a food item occurs as a percent of total number of occurrences for all food items (Kelly 1991, Wagner 1993). Percent of occurrence data is presented for all food items (Fig. 1), all vegetative food items (Fig. 2), and all mammalian food items (Fig. 3). We grouped some food items due to difficulties in differentiating among species and combined items occurring at low frequencies into an “other” category. Results We analyzed 415 scats to evaluate coyote food habits between May 2005 and July 2006 (Table 1). No single food item occurred in all 15 months sampled. White-tailed deer, lagomorphs, and insects, primarily Orthoptera, occurred in 14 months each, and small mammals occurred in 13 months (Table 1). A single food item made up greater than 90% of the scat volume in 227 (54.7%) of the samples we collected. The coyote diet was dominated by plant matter from May 2005 through November 2005 as well as from June through July 2006 (Fig. 1). The occurrence of individual plant food items consumed varied with expected availability based on known fruiting times of these species (Fig. 2). Prunus spp. Linnaeus (wild 138 Southeastern Naturalist Vol.7, No. 1 Table 1. Percent of scats containing common coyote food items at the Savannah River Site, SC by month from May 2005 through July 2006. 2005A 2006A May June July Aug Sept Oct Nov Dec Jan Feb Mar Apr May June July Food item (13) (53) (22) (51) (32) (26) (26) (25) (18) (27) (30) (13) (39) (31) (9) Plants Arachis hypogaea 4 17 4 3 8 3 Carya spp. (nuts) 8 4 15 13 3 Crataegus spp. 25 13 15 4 8 6 Diospyros virginiana 18 78 81 31 8 Phytolacca americana 5 51 59 77 62 36 17 4 3 Poaceae (leaves) 4 6 4 3 8 3 10 Prunus serotina 4 59 10 6 4 22 Prunus spp. 85 43 9 6 54 52 33 Quercus spp. (acorns) 4 8 11 7 Rubus spp. 23 64 27 2 8 26 58 89 Vaccinium spp. 18 37 4 4 6 Vitis spp. 30 25 12 OtherB 5 6 4 8 Animals Aves 2 2 4 4 6 4 10 18 6 11 Castor canadensis 2 6 3 11 Dasypus novemcinctus 5 4 4 4 8 5 Neotoma fl oridana 2 11 3 23 5 Odocoileus virginianus (adult) 15 8 14 6 3 15 40 11 23 37 15 5 10 11 Odocoileus virginianus (fawn) 31 15 18 2 38 23 Orthoptera 8 21 27 31 31 31 12 8 11 3 31 31 32 44 Procyon lotor 6 3 4 4 7 8 3 Sciurus spp. 15 3 6 3 4 4 8 6 4 10 8 Small mammal 8 8 5 10 9 4 8 11 15 10 23 3 10 Sus scrofa 11 14 10 4 4 6 31 10 8 8 3 Sylvilagus spp. 8 6 9 6 16 4 4 12 11 31 17 8 5 3 OtherC 3 8 8 6 4 7 8 ANumbers in parentheses are sample sizes. BIncludes Ilex decidua, Ostrya virginiana, Passifl ora incarnata, and Pyrus sp. CIncludes Canis latrans, Didelphis virginiana, Lynx rufus, Ondatra zibethicus, Urocyon cinereoargenteus, and egg fragments. 2008 J.D. Schrecengost, J.C. Kilgo, D. Mallard, H.S. Ray, and K.V. Miller 139 plums) occurred in 85% of scats in May 2005 and 54% of scats in May 2006. Rubus spp. Linnaeus (blackberries) occurred in 64% of scats in June 2005, 27% of scats in July 2005, 58% of scats in June 2006, and 89% of scats in July 2006. Prunus serotina Ehrhart (black cherry) occurred in 59% of scats during July 2005. Phytolacca americana Linnaeus (pokeberry) occurred in 51% of scats during August 2005 and 62% of scats during November 2005. Diospyros virginiana Linnaeus (persimmon) occurred in 78% of scats during September 2005 and 81% of scats during October 2005. Figure 1. Monthly percent of occurrence for 4 major groups of food items in coyote scats collected on the Savannah River Site, SC, May 2005 through July 2006. Figure 2. Monthly percent of occurrence for major vegetative food items in coyote scats collected on the Savannah River Site, SC, May 2005 through July 2006. 140 Southeastern Naturalist Vol.7, No. 1 During December through April, the coyote diet was dominated by animal food items (Fig. 1), although percent occurrence of individual mammalian food items varied throughout the year (Fig. 3). White-tailed deer occurred in 40% of scats during December and 37% of scats during March. Conception dates of white-tailed deer at SRS (Rhodes et al. 1991) plus a 200-day gestation period (Haugen 1959, Verme 1965) place parturition primarily in the months of May and June. During 2005, white-tailed deer fawns were present in scat collected in May (31%), June (15%), July (18%), and August (2%). During 2006, fawns were present in scat collected in May (38%) and June (23%). Wild hogs and Sylvilagus spp. (rabbits) each occurred in 31% of scats during February. Orthoptera occurred in 31% of scats during April (Table 1). Discussion From spring through late fall, coyotes at SRS fed heavily on soft mast when it was available. However, from December through March, when vegetative food items were less abundant, coyotes relied heavily on mammalian food items. Small- and medium-sized mammals may also be more vulnerable at this time of year due to a decrease in herbaceous cover. Orthopterans, primarily grasshoppers, were common in scats throughout the year, but they rarely comprised a significant portion of the volume of the scats. Similarly, birds were consumed throughout the year, but at a relatively low percent of occurrence. Greatest bird consumption occurred during May 2006 (18%). Although this timing was likely coincidental with avian nesting activity and egg shell fragments were observed in scat, low occurrence during 2005 suggests large annual fl uctuations in bird consumption. Small mammals and lagomorphs appear to be less important in the diets of coyotes at the SRS than elsewhere. Other studies have reported high Figure 3. Monthly percent of occurrence for major mammalian food items in coyote scats collected on the Savannah River Site, SC, May 2005 through July 2006. 2008 J.D. Schrecengost, J.C. Kilgo, D. Mallard, H.S. Ray, and K.V. Miller 141 occurrences of lagomorphs (Bartel and Knowlton 2005, Thornton et al. 2004) and small mammals (Bartel and Knowlton 2005, Hall 1979, Smith and Kennedy 1983) in coyote diets. For example, Blanton and Hill (1989) reported 36.5% frequency of occurrence of Sylvilagus spp. in 523 coyote scats and 9 coyote stomachs collected during summer in Mississippi, Alabama, Kentucky, and Tennessee. Bowyer et al. (1983) reported 28.2% annual frequency of occurrence for Rodentia in coyote scats in California. In Louisiana, Wilson (1967) found rodent remains occurring in 75% of coyote stomachs during spring and fall and lagomorph remains occurring in 80% of coyote stomachs during summer. Although both of these groups were present in the diet of coyotes at SRS throughout most of the year, their importance was limited compared to other food items, and occurrence was greatest during winter and early spring (Table 1). Specifically, the highest occurrence of small mammals in coyote scats on SRS occurred during April 2006 (23%), and the highest occurrence of lagomorphs occurred during February 2006 (31%). Although these peak values approach the annual or seasonal occurrences reported in other studies, they were limited to single months. Neither group occurred in >17% of scats collected during any other months (Table 1). Our data does suggest that lagomorphs and small mammals were important relative to other mammalian food items during September and October of 2005 (Fig. 3). However, during this time, the coyote diet was dominated by vegetative food items, and mammalian food items made up <15% of the food items consumed (Fig. 1). Occurrence of wild hog in scats was greatest during February (31%), corresponding with the peak of hog control (January through March), during which 69 hog carcasses were disposed of on SRS. Similarly, Wagner (1993) found hog remains in 17–25% of scats in Bolivar County, MS and reported peak occurrence coincident with the peak in sport hunting of hogs. Hog remains in scats occasionally contained small hooves, suggesting predation on piglets, but most hog occurrence in scats consisted of the large, coarse hairs of adult hogs. Coyotes undoubtedly scavenge adult hog carrion, but we are aware of no published reports of coyote predation on adult hogs. Nevertheless, we cannot eliminate the possibility of predation on adult hogs, particularly given the somewhat atypical use of other food items by SRS coyotes. Fruit use by coyotes has been documented by numerous studies (Andelt et al. 1987, Blanton and Hill 1989, Murie 1951, Thornton et al. 2004, Wooding 1984). However, the timing of fruit availability and species use is variable. Wooding (1984) reported persimmons in 67% of scats and stomachs during fall in Mississippi and Alabama, similar to the findings of this study during September through November (31–81%). Blackberry and pokeberry have also been reported at significant levels in several southeastern states (Blanton 1988). However, high occurrence of wild plums in the diet of coyotes has not been previously reported. During the months of May and June, wild plum occurred in 43–85% of the coyote scats collected on the SRS during 2005 and 2006. The differences in occurrence of soft mast within the same region further emphasize the need for locally specific coyote food-habit studies. White-tailed deer were consumed in every month except October. Roadkilled deer carcasses are available to coyotes in limited quantities throughout 142 Southeastern Naturalist Vol.7, No. 1 the year. During 2005, 72 road-killed deer were recovered along the SRS roadways (P.E. Johns, Savannah River Ecology Laboratory, Aiken, SC, unpubl. data) and most were disposed of in wooded areas on site. This explains a portion of the deer remains encountered in coyote scats because we often observed scavenging by coyotes at monitored carcasses. The absence of white-tailed deer in October is coincident with the increase in seasonal soft mast consumption as persimmon and pokeberry fruits became available. Increased use of whitetailed deer during November and December corresponded with the timing of controlled hunts on SRS. Apparently coyotes consumed deer crippled or un-recovered during hunts. We believe it is unlikely that hunting pressure made deer more susceptible to predation. D’Angelo et al. (2003) found that female whitetailed deer on SRS resumed normal movement patterns within 13 hours after hunt-associated disturbances, suggesting that risk of coyote predation likely would not be greatly increased by hunting activity. White-tailed deer remains also occurred in a large proportion of scats during March 2006 (Table 1). Increased coyote use of deer during late winter has been associated with increased deer vulnerability due to winter severity and deep snows (Patterson et al. 1998). However, winters are mild in South Carolina. Deer-vehicle collision data from the SRS suggests there was no significant increase in availability of deer carrion at this time. Although coyotes may have been killing adult deer, it is more likely that coyotes were returning more often and from greater distances to the available carrion and consuming even portions of the deer hides, and thus hair. This explanation could account for the increased occurrence of deer remains in scats during March. White-tailed deer fawns were an important component of the diet, occurring in 15–38% of scats during May through July, coincident with the timing of parturition and fawn rearing. During this time, fawns comprised the largest proportion of mammalian food items encountered, although blackberries and wild plums were the most frequently encountered items in scats overall (Table 1). Similarly, Blanton and Hill (1989) reported 31% average occurrence of deer in the summer coyote diet across seven southeastern study areas. Data from Mississippi and Alabama (Wooding 1984) and from Louisiana (Hall 1979) are comparable. Blanton and Hill (1989) observed a decreased use in other food items as fawns became available in areas of high deer density, suggesting that coyotes may select fawns over less profitable food items. Blanton and Hill (1989) used an annual deer-harvest density of 1 per 48 ha or greater to define high-density populations. Deer-harvest density on the SRS was 1 per 372 ha during 2005. On SRS, fawns may be more profitable than alternative food sources despite low deer density (estimated deer density of 1 per 26 to 29 ha). Together with the high occurrence of fawn remains in coyote scats reported herein, several lines of evidence suggest the possibility that coyotes may be impacting deer recruitment at the SRS. These include high coyote-induced mortality among radio-collared fawns during a 2006–2007 study, a 54% decline in the proportion of fawns/doe in the harvest between the periods 1965–1993 and 1997–2006, and a 45% decline in an annual spotlight index from 1991–2007 (J.C. Kilgo and H.S. Ray, unpubl. data). In addition, the increased use of mammalian food items when soft mast availability is limited, suggests that a spring 2008 J.D. Schrecengost, J.C. Kilgo, D. Mallard, H.S. Ray, and K.V. Miller 143 soft-mast failure could result in increased predation of white-tailed deer fawns (Andelt et al. 1987). Our data presents further evidence in support of this theory because increases in mammalian food-item use were correlated with decreases in seasonal soft-mast availability. Additional research is warranted to better understand the potential impacts of the recently established coyote on whitetailed deer and other species in southeastern ecosystems. Acknowledgments We thank M. Schrecengost for dedicated assistance in data collection. We acknowledge the support of D. Osborn and P.E. Johns and the cooperation of US Forest Service-Savannah River employees. I.L. Brisbin, J.J. Mayer, and an anonymous reviewer provided helpful comments on the manuscript. 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