Regular issues
Special Issues

Southeastern Naturalist
    SENA Home
    Range and Scope
    Board of Editors
    Editorial Workflow
    Publication Charges

Other EH Journals
    Northeastern Naturalist
    Caribbean Naturalist
    Urban Naturalist
    Eastern Paleontologist
    Eastern Biologist
    Journal of the North Atlantic

EH Natural History Home

Local-Scale Difference of Coyote Food Habits on Two South Carolina Islands
Cady R. Etheredge, Sloane E. Wiggers, Olivia E. Souther, Lindi L. Lagman, Greg Yarrow, and Jamie Dozier

Southeastern Naturalist, Volume 14, Issue 2 (2015): 281–292

Full-text pdf (Accessible only to subscribers.To subscribe click here.)


Site by Bennett Web & Design Co.
Southeastern Naturalist 281 C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 22001155 SOUTHEASTERN NATURALIST 1V4o(2l.) :1248,1 N–2o9. 22 Local-Scale Difference of Coyote Food Habits on Two South Carolina Islands Cady R. Etheredge1,*, Sloane E. Wiggers2, Olivia E. Souther2, Lindi L. Lagman3, Greg Yarrow3, and Jamie Dozier4 Abstract - Canis latrans (Coyote) is regarded as a classic generalist predator that has recently established large populations throughout the southeastern US. To better understand how Coyote food habits in the Southeast may differ on an extremely small spatial scale, we collected a total of 305 Coyote scats from 2009 to 2011 on 2 islands separated by a 1.4–2.5-km-wide expanse of low saltwater-marsh on the coast of Georgetown, SC. We identified diagnostic remains of prey items to the lowest possible taxonomic level. A multiresponse permutation procedure revealed differences in Coyote diet composition between islands (A = 0.0090, P < 0.0001). Subsequent indicator-species analysis revealed a total of 4 food items that served to differentiate diet between islands: birds, Sus scrofa (Wild Hog), Ilex sp. (holly) fruit, and lagomorphs. Our results demonstrate that Coyote food habits and their potential ecosystem effects may vary widely on a very local scale. This finding may be of particular concern to biologists attempting to utilize published diet studies to inform Coyote management strategies. Our study also documented some of the highest levels of bird consumption by Coyotes published to date; we detected bird remains found in 42.45– 59.80% of scats. Introduction Canis latrans Say (Coyote) is a new invader of ecosystems across the southeastern US (Parker 1995) and could have potentially large impacts on population dynamics of prey in the region (Kilgo et al. 2012). Although the basic ecology of Coyotes has been widely studied in the western US, these studies may be of limited use in understanding specific food habits of southeastern populations because of the eastern Coyote’s larger body size and the extreme behavioral plasticity (Schrecengost et al. 2008). Like western populations, diets of southeastern Coyotes are comprised largely of rodents, vegetation, and lagomorphs, with the abundance of items such as fruit, domestic animals, livestock, commercial crops, wild ungulates, and birds varying greatly based on prey availability (Parker 1995). A number of authors have addressed Coyote diet in regions of the Southeast where populations have been established since the 1930s, but diet investigations are lacking throughout Georgia and the Carolinas where Coyote populations are still expanding 1Division of Wildlife Ecology, University of Wisconsin at Stevens Point, Stevens Point, WI 54481. 2Department of Biological Sciences, Clemson University, 132 Long Hall, Clemson, SC 29634. 3School of Agricultural, Forest, and Environmental Sciences, Clemson University, 251 Lehotsky Hall, Clemson, SC 29634. 4South Carolina Department of Natural Resources, 1 Yawkey Way South, Georgetown, SC 29440. *Corresponding author - Manuscript Editor: Michael Conner Southeastern Naturalist C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 282 (Schrecengost et al. 2008). Detailed studies of localized food habits are of vital importance to wildlife biologists throughout the region, who base their management decisions on the best available information (Smith and Kennedy 1983). Studies investigating differences in the diet of generalist species living in different areas should compare data from 2 or more study areas with minimal connectivity between them to ensure items consumed in one study area are not deposited in another. Animal movement between areas utilized in such studies may be limited by distance between sites or by some barrier to movement between study areas that limits connectivity. Often, studies investigating the relationship between diet and habitat are coupled with radio-telemetry studies of space use, in which animal groups are known to occupy defined areas and movement between areas has been shown to be limited (e.g., Lander et al. 2009, Lavin et al. 2003, Lea et al. 2008, Morey et al. 2007). Researchers investigating differences in diet without radio-telemetry often use reported home ranges of the target species as a physical-distance proxy for information about movement. For example, Farias and Kittlein (2008) chose sites separated by 15 km to test for differences in the diet of Lycalopex gymnocercus Fischer (Pampas Fox), which have an average home-range size of 0.45 km2. Utilizing information on average home-range size in this manner only takes into account physical distance between sites. However, areas that are physically close together but have low connectivity between patches due to natural or anthropogenic factors should allow for similar comparisons between groups. The goal of our study was to investigate food habits of southeastern Coyotes in an island system. Our objectives were to (1) document Coyote diet on 2 islands on the coast of South Carolina, (2) test for differences in Coyote food habits between areas in close proximity but with potentially low connectivity between areas, and (3) speculate on what these potential differences in Coyote diet might mean for wildlife managers in the southeastern US. Field-Site Description The Tom Yawkey Wildlife Center Heritage Preserve (TYWCHP) is a 9700-ha wildlife preserve off the coast of Georgetown, SC. The TYWCHP consists of Cat, South, and North Islands (Fig. 1); Cat and South Islands were the focus of this study. Cat Island is separated from the mainland by the Atlantic Intracoastal Waterway and contains Pinus palustris Mill. (Longleaf Pine) flatwoods, freshwater bogs, saltwater and freshwater waterfowl impoundments, and planted wildlife openings. Pine flatwoods are burned on a 2-year rotation to prevent hardwood intrusion (Dozier 1996). Upland areas on Cat Island include a wide variety of dominant plant species, including Quercus marilandica Muenchh. (Blackjack Oak), Pteridium aquilinum L. (Bracken Fern), Vaccinium spp. (blueberries) and Gaylussacia spp. (huckleberries). South Island consists mainly of saltwater waterfowl impoundments, maritime forest, and barrier beach with Winyah Bay to the north and the Atlantic Ocean to the east. Upland areas on South Island include mainly maritime forest communities dominated by Quercus virginiana Mill. (Southern Live Oak), Ilex vomitoria Sol. (Yaupon), Juniperus virginiana L. (Eastern Red Cedar), Magnolia grandiflora L. Southeastern Naturalist 283 C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 (Southern Magnolia), Pinus taeda L. (Loblolly Pine), and Sabal minor Pers. (Dwarf Palmetto). Cat Island is roughly 3 times as large as South Island (4525 ha and 1507 ha, respectively), but South Island includes a larger area of managed wetlands (485 ha and 702 ha on Cat Island and South Island, respectively; J. Dozier, unpubl. data). The TYWCHP is recognized as a western hemispheric shorebird preserve and an Audubon Important Bird Area due to the large numbers of waterfowl, shorebirds, and wading birds that utilize its managed wetlands (Hopkins-Murphy 1989). Figure 1. Cat Island and South Island on the Tom Yawkey Wildlife Center and Heritage Preserve, Georgetown, SC. Southeastern Naturalist C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 284 The TYWCHP is managed by the South Carolina Department of Natural Resources (SCDNR) and is closed to hunting and to general public access. Both islands contain a variety of mammals, including a large number of small-mammal species (<200 g), Sylvilagus floridanus J.A. Allen (Eastern Cottontail), Didelphis virginiana Kerr (Virginia Opossum), Procyon lotor L. (Raccoon), and Lynx rufus Schreber (Bobcat) (J. Dozier, pers. observ.). Odocoileus virginianus Zimmermann (White-tailed Deer), Sus scrofa L. (Wild Hog), and Sciurus carolinensis Gemlin (Eastern Grey Squirrel) occupy both islands but are more commonly seen on Cat Island (J. Dozier, pers. observ.). Sciurus niger L. (Fox Squirrel) is found primarily on Cat Island (J. Dozier, pers. observ.). White-tailed Deer density on both islands averages ~1 deer/8 ha in upland areas (Dozier 1996). SCDNR staff removed 10– 16 White-tailed Deer and ~20 Wild Hogs from Cat Island during each year of the study period; the carcasses generated by these management activities were deposited uncovered at a disposal area on South Island where Coyotes had access to the carrion. The first report of a Coyote on the TYWCHP was from Cat Island in 2006 (J. Dozier, unpubl. data). South Island is separated from Cat Island by a 1.4–2.5-km-wide expanse of low saltwater marsh. The marsh is tidally influenced and exposed at low tide. It is characterized by dense stands of Spartina alterniflora Loisel. (Salt Marsh Cordgrass) and Juncus roemerianus Scheele (Black-needle Rush) with thick layers of organic matter and silt. The 2 islands are connected by a 3.2-km-long causeway, which is the only road for vehicular traffic between the islands. Coyotes seen travelling between islands on the causeway are targeted by SCDNR staff; 1 Coyote was shot on the causeway and 1 was hit by a truck during the study period. Methods We collected Coyote scats on transects along roads, dikes, and through beachfront dunes from May 2009 to July 2009 and January 2010 to December 2010. We established 7 transects on both Cat Island and South Island; all transects were ~2–3 km long. We traveled each transect by foot, bicycle, or truck at least twice during each season, with seasons defined as winter (December–February), spring (March–May), summer (June–August), and fall (September–November). We also collected scats opportunistically during the course of other field work and stored them in plastic bags at room temperature before processing. We hand-washed each scat with water over a 1-mm-mesh screen and air dried them. We removed diagnostic remains of diet items (e.g., dorsal guard hairs, bones, teeth, claws, seeds) from scats and identified the items to the lowest taxonomic level possible using reference collections at the Campbell Museum of Natural History, Clemson, SC, and with identification keys (Martin and Barkley 1961, Moore et al. 1974, Roest 1986). We distinguished Coyote and Bobcat scats by size and shape (Murie and Elbroch 2005). Plant matter deemed to have been collected incidentally with the sample (oak leaves, pine needles) and not likely purposefully ingested by a Coyote (grass, seeds) was removed from analysis, as were intact, undigested insects that may have been feeding on collected samples. Southeastern Naturalist 285 C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 We calculated the proportion of each diet item utilized as the percent of scats (the number of scats with a diet item x 100 / total number of scats). Shannon’s diversity index was calculated for each island to provide a measure of diet diversity, after which diet items found in <1% of scats for both islands combined were eliminated from further analysis. Multi-response permutation procedures (MRPP; Mielke and Berry 2001) were performed with a Sorenson (Bray-Curtis) distance measure to test the hypothesis of no difference in Coyote diet composition between islands. MRPP is a multivariate technique that calculates a distance matrix to compare the overall composition of 2 groups (here, islands), to test the null hyposthesis of no difference between groups. Subsequently, we used pairwise chi-squared tests to test for differences between islands for each food item. We also conducted an indicator-species analysis (ISA) to assess whether different diet items differentiated between islands (Dufrene and Legendre 1997). ISA is a multivariate method that determines which species are more closely identified with one group (island) than another. ISA utilizes both the species abundance and frequency to calculate an indicator value (IV) from 0–100 for each species (McCune et al. 2002). A value of 0 in this application suggested that a species was completely absent from an island, while a value of 100 signified that a species was found in every scat taken from that island (but not from the other). We tested for significance of the ISA with a Monte Carlo test with 4999 permutations. Shannon’s diversity indices, MRPP, and ISA were conducted with PC-ORD (MJM Software Design, Gleneden Beach, OR); SAS (SAS Institute, Cary, NC) was used for chi-squared tests. We set a significance level of α = 0.05 for all tests. We did not collect relatively small scats (<2.5 cm in diameter) which might have been confused with Bobcat scat (Grigione et al. 2011). These smaller scats located in the dunes or exposed areas of South Island were often observed for >3 months after initial observation, suggesting that it might be difficult to determine when larger (Coyote) scats were actually deposited. For this reason, and because low sample sizes precluded seasonal or annual comparisons between islands, we made no attempt to distinguish between seasonal or annual differences in diet. Results We collected a total of 106 and 199 scats on Cat Island and South Island, respectively, with more scats collected on South Island in 5 of 7 sampling periods (Table 1). We identified 44 total items on both islands combined—32 species from Cat Island scats and 39 from South Island. Shannon’s diversity indices were similar for each island (Cat Island = 2.44, South Island = 2.46), but MRPP showed a significant difference between Coyote diets on the 2 islands (A = 0.0090, P < 0.0001). Table 1. Coyote scats collected by season on Cat Island and South Island at the Tom Yawkey Wildlife Center and Heritage Preserve, Georgetown, SC. Spring Summer Winter Spring Summer Fall Winter 2009 2009 2010 2010 2010 2010 2010–2011 Total Cat Island 4 11 36 32 3 15 5 106 South Island 34 23 50 44 31 13 4 199 Southeastern Naturalist C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 286 Sigmodon spp. (cotton rats) were the most common food item found in Cat Island scats, followed by birds, vegetation, and Peromyscus spp. (deer mice). Birds were the most common item found in South Island samples, followed by cotton rats, vegetation, and Neotoma spp. (wood rats; Table 2). Cat Island samples comprised a larger percent of scats containing Wild Hog, lagomorphs, Diospyros sp. (persimmon), and soricomorphs, while South Island samples contained more birds, crabs, Mephitis mephitis Shreber (Striped Skunk), and mustelids (Table 2). ISA yielded significant indicator values for 3 animal groups and 1 plant genus: birds, lagomorphs, Wild Hogs, and Ilex spp. (Table 2). Table 2. Percent of scats, item rank, and indicator value for diet items found in Coyote scats on Cat Island and South Island at the Tom Yawkey Wildlife Center and Heritage Preserve, Georgetown, SC, 2009–2011. Percent of scats was calculated as the number of scats with a diet item (n) x 100 / total number of scats (N). Indicator values are the observed maximum indicator value for both islands, and IV P-values are the result of a Monte Carlo test of significance based on 4999 randomizations. * indicates a significant difference in percent of scats between islands (chi-square test; P < 0.05). Cat Island (N = 106) South Island (N = 199) Indicator Diet item % (n) Rank % (n) Rank value (IV) P Small mammals Microtus sp. 18.87 (20) 6 13.07 (26) 6 11.1 0.1928 Neotoma sp. 11.32 (12) 10 18.59 (37) 4 12.4 0.1398 Oryzomys sp. 7.55 (8) 11 7.04 (14) 11 3.9 0.1398 Peromyscus sp. 23.58 (25) 4 17.09 (34) 5 13.6 0.2222 Rattus sp. 0.66 (2) 15 3.52 (7) 15 2.4 0.4861 Scuridae 2.83 (3) 14 4.02 (8) 14 2.5 0.7540 Sigmodon sp. 54.72 (58) 1 46.73 (93) 2 29.2 0.2899 Soricidae* 13.21 (14) 8 9.05 (18) 9 7.8 0.3263 Midsized herbivores Lagomorpha* 21.70 (23) 5 10.05 (20) 8 14.8 0.0050 Large herbivores Odocoileus virginianus 3.77 (4) 13 6.03 (12) 12 4.0 0.4323 Sus scrofa* 14.15 (15) 7 3.02 (6) 16 11.7 0.0018 Mesopredators Didelphis virginiana 12.26 (13) 9 7.54 (15) 10 7.5 0.2178 Mephitis mephitis* 0.33 (1) 16 1.51 (3) 18 1.0 1.0000 Mustelidae* 0.00 (0) 2.51 (5) 17 2.6 0.1622 Procyon lotor 1.89 (2) 15 1.01 (2) 19 1.2 0.6179 Other Aves* 42.45 (45) 2 59.80 (119) 1 37.5 0.0038 Decopoda 1.89 (2) 15 2.51 (5) 17 1.5 1.0000 Diospyros sp.* 2.83 (3) 14 0.50 (1) 20 1.5 0.1322 Ilex sp.* 0.94 (1) 16 10.55 (21) 7 5.2 0.0016 Insecta 7.55 (8) 11 3.52 (7) 15 3.8 0.1660 Reptilia 4.72 (5) 12 3.02 (6) 16 2.9 0.5333 Uknown seeds 7.55 (8) 11 5.03 (10) 13 4.5 0.4663 Vegetation 32.08 (34) 3 31.66 (63) 3 16.9 0.9064 Southeastern Naturalist 287 C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 Discussion As expected with a generalist predator, differences in prey availability and habitat type may explain the differences in Coyote diet found in this study (Dumond and Villard 2001, Morey et al. 2007). Birds, lagomorphs, Ilex sp., and Wild Hogs were identified by ISA as being important contributors of overall differences in food habits between the islands. South Island includes a larger area of managed wetlands (702 ha, ~47% of the overall area; J. Dozier, unpubl. data) that support more wading birds and shorebirds than Cat Island (J. Dozier, Christmas Bird Count, unpubl. data). Upland areas of South Island are also dominated by Yaupon, which is commonly found in the diets of mammalian generalists (Miller and Miller 2005). Comparatively more wading bird and shorebird habitat and an abundance of Yaupon on South Island could explain a greater percent of scats with bird remains and Yaupon seeds in scat samples from that island. Likewise, Wild Hog populations are well established on Cat Island, but not South Island (J. Dozier, unpubl. data), and more Cat Island scats contained Wild Hog remains. Although no data on the distribution and abundance of lagomorphs exist for the TYWCHP, Cat Island has more upland habitat than South Island, most of which is comprised of Longleaf Pine flatwoods with a diverse herbaceous understory that should favor lagomorphs (Yarrow and Yarrow 1999). Thus, differences in habitat structure between the 2 islands could explain the presence of significantly more lagomorphs in Cat Island samples. Pair-wise chi-squared tests detected 4 additional diet items from scats that showed a significant between-island difference. However, none of these were identified by ISA as important drivers of overall diet. Crab, Striped skunk, and mustelid items were all found more often in South Island samples, but likely not in quantities large enough to influence overall diet (<3% on both islands). We also found Soricomorphs more commonly in Cat Island than South Island samples (13.21% and 9.05% on Cat Island and South Island, respectively) and more commonly overall than the other 3 of these additional diet items, but perhaps not commonly enough to be included in ISA. Other southeastern Coyote food-habit studies have documented rodents and vegetation as major diet items (e.g., Crimmins et al. 2012, Grigione et al. 2011, Hall 1979, Smith and Kennedy 1983), but no other southeastern studies to date have documented such a large avian component in Coyote diets. No other study in the Southeast has documented more than 20% of Coyote scat or stomach samples containing bird remains (e.g. Crimmins et al. 2012, Grigione et al. 2011, Schrecengost et al. 2008, Smith and Kennedy 1983), but in our study, 42–60% of scats on the TYWCHP contained feathers or bird bones. Several researchers have documented low levels of Coyote consumption of songbirds, most of which are listed as unidentified passeriformes (Gipson 1974, Hall 1979, Hoerath 1990, Michaelson 1975). However, Hall (1979) was able to identify 10 different songbird species from recovered flight feathers in Louisiana Coyote scats. We did not recover flight feathers during the present study. Instead, most of the feathers in our colleceted scats were downy white or gray, and lacked any identifiable markings (Scott and McFarland 2010). While a lack of distinguishing marks makes it difficult to determine which bird Southeastern Naturalist C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 288 species or groups the Coyotes at our sites were utilizing, it appears that during this study wading birds may have been more likely to be preyed upon compared to other avian taxa (e.g. passerines). Given the relatively larger body size of wading birds than passerines, Coyotes at our sites may well have selectively consumed the bodies of wading birds, avoiding the wings and larger feathers that would have aided in species identification. In contrast, if Coyotes had consumed passerines, which may be easier to consume as whole birds, we would likely have found flight feathers and other identifiable remains in the scat. Coyote consumption of wading birds at this study site is also more likely given the large numbers of wading birds that utilized waterfowl impoundments on the TWYCHP each year. Future studies of Coyote diet in coastal areas may be able to utilize stable isotope techniques to distinguish between songbirds— which typically consume terrestrial insects, fruits and seeds—and wading birds—which utilize aquatic prey (e.g., Hilderbrand et al. 1996). The between-site differences in Coyote diet demonstrated in this study might also provide a basis for hypotheses about Coyote movements between islands. If Coyotes moved readily across the marsh between islands, it follows that there would likely be enough items consumed on one island and deposited on the other to obscure any habitat-driven differences between areas. For this reason, animal movement between study areas is often a major concern for studies attempting to document differences in food habits between areas (e.g., Lander et al. 2009, Lavin et al. 2003, Lea et al. 2008, Morey et al. 2007). However, no home-range studies of Coyotes have been published on the TYWCHP to date, so for our study, the interpretation of food habits to understand movements depends heavily on the distance Coyotes travel after consuming a food item before leaving scat. While this distance has not been estimated for Coyotes, Marucco et al. (2008) found that Canis lupus L. (Gray Wolf) may travel up to 2 days before depositing scat after a kill. Dumond and Villard (2001) used a radius of 5 km around Coyote scat samples to denote where ingestion of the food item likely occurred. If we assumed a 5-km radius for our study at TYWCHP, there would be a large amount of overlap between islands, potentially indicating that in the absence of the marsh separating the islands, Coyotes should be equally likely to deposit prey remains in scat on either island. Long-distance movements of Coyotes (>15 km/day) have been reported in the Southeast but are usually attributed to transient animals (Hinton et al. 2012). Further, different diet items may have greater utility in providing information on animal movements. In particular, fruits may not be as good of an indicator of Coyote movements at our sites because they are processed and expelled more quickly than mammalian food items (Andelt and Andelt 1984). Although ISA detected a difference in 1 fruit species (Yaupon), we also found important between-island differences in 3 vertebrate food items (lagomorphs, birds, and Wild Hogs). Taken together, our data could suggest limited movement of Coyotes between islands. Further, understanding food habits in light of animal movements in this manner could suggest an exciting new line of research regarding generalist predators. Despite equal search effort on both islands, we observed more scats on South Island than Cat Island. This difference could have been due to a greater density of Southeastern Naturalist 289 C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 Coyotes on South Island than Cat Island, increased persistence of scats on South Island than those on Cat Island, or increased detectability of scats on South Island than those on Cat Island. No population estimates have been made for Coyotes on the TYWCHP and hence, a comparison of density between islands is not available. However, anecdotal evidence suggests that there is increased persistence of scats on South Island, where scats are more exposed in open habitats on beach dunes and along dikes. Scat persistence has been addressed in a variety of different environments (e.g., Godbois et al. 2005, Livingston et al. 2005, Sanchez et al. 2004), but no studies to date have documented scat persistence on coastal habitats in the Southeast. Scats not collected in our study that were located in open areas exposed to the sun were often observed to last >3 months in the field, perhaps due to quick desiccation. We did not assess potential differences in Coyote diet related to season because of the uncertainty of when scats were actually deposited (as opposed to collected). Because more scats were collected on South Island in the spring and summer, seasonal variability could be influencing our detection of differences in island diets. Further, if more scat was found on South Island during the spring and summer seasons because of a shift in Coyote space use from one island to the other, differences in diet between islands could be driven by Coyote movement, and not by habitat differences between islands. Scats may have been more detectable on South Island than Cat Island because of the more exposed nature of transects on South Island. Diet studies of generalist carnivores that rely on identification of items from scat often suffer from biases related to various consumption patterns and assimilation efficiencies of different groups of food items (Andelt and Andelt 1984, Marucco et al. 2008, Rühe et al. 2008). For example, carnivores that utilize carcasses of large mammals may consume more meat or organs and less hair or bones than those that consume whole rodents, potentially causing the importance of large mammals in carnivore diets to be underrepresented because remains from meat or organs are less likely to appear in scat compared to hair samples or bone fragments (Marucco et al. 2008). Similarly, we found no egg shells (either avian or reptilian) in this study, despite Coyotes on the TYWCHP being the main predator of Caretta caretta L. (Loggerhead Sea Turtle) nests on South Island (Eskew 2012). Coyotes on South Island break open turtle eggs on the beach and lick out the yolk, which leaves no diagnostic remains in scat (C.R. Etheredge, pers. observ.). Even so, other studies have found egg shells in Coyote scat (avian: Litvaitis and Shaw 1980, Wagner and Hill 1994; reptilian: Wooding et al. 1984). Further, no remains of Loggerhead Sea Turtle hatchlings were documented in the present study, even though Coyotes are a known predator of hatchlings on South Island (Eskew 2012), suggesting that studies of Coyote diet based on scat sampling may be inadequate to detect the potential for Coyote impacts on some species of special conservation concern. Coyote impacts on White-tailed Deer are particularly concerning for southeastern wildlife managers, and several authors have documented large proportions of White-tailed Deer remains in Coyote scats and stomachs (Blanton and Hill 1989, Crimmins et al. 2012, Schrecengost et al. 2008). These findings have suggested Southeastern Naturalist C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 290 that Coyote depredation, particularly on fawns, may have a profound region-wide effect on White-tailed Deer populations (Kilgo et al. 2012), especially in areas where White-tailed Deer densities are very high (Blanton and Hill 1989). The present study found relatively low frequencies of White-tailed Deer (less than 7% of scats on both islands) compared to other studies (Schrecengost et al. 2008), which may be a function of lower White-tailed Deer densities on the TYWCHP (Blanton and Hill 1989). It is also possible that we were less likely to collect scats containing Whitetailed Deer remains if Coyote consumption of larger prey items influenced the rate of scat decomposition, as was documented for Bobcats in Georgia by Godbois et al. (2005). We did not attempt to differentiate between adult White-tailed Deer and fawns, although it is worth noting that Coyotes readily scavenged White-tailed Deer left uncovered at the carcass-disposal area on South Island (C.R. Etheredge, pers. observ.). Differences in food habits documented at the local scale employed in this study suggest that diet may differ within a region, and hence that regional generalizations about Coyote diets may be misleading. Landowners and wildlife managers alike should understand that even a study conducted in the same state or county likely does not necessarily reflect conditions on their own property. Studies testing predictive hypotheses about Coyote food habits based on habitat types or prey-population sizes across the Southeast (e.g., Blanton and Hill 1989) will likely be more useful to managers than smaller-scale studies reporting variation in Coyote diet based on season, habitat, or prey availability which has already been well established for Coyotes. Acknowledgments We offer extreme gratitude to S. Miller, A. Kremenski, and many others for assistance in the field and in the lab. We also thank R. Baldwin, P. Jodice, J. Armstrong, P. Gerard, and others for comments on the manuscript and S. Miller and N. Etheredge for assistance with reference collections. C. Marion was helpful with data analysis. C.R. Etheredge was supported by a SCDNR State Wildlife grant and a Marion E. Bailey Assistantship. The Yawkey Foundation, the Clemson University South Carolina Life program, and the Clemson University Creative Inquiry program provided funding and logistical support for this project. Literature Cited Andelt, W.F., and S.H. Andelt. 1984. Diet bias in scat deposition-rate surveys of Coyote density. Wildlife Society Bulletin 12:74–77. Blanton, K., and E. Hill. 1989. Coyote use of White-tailed Deer fawns in relation to deer density. Proceedings of the Annual Conference of the Southeastern Association of Fish and Wildlife Agencies 43:470–478. 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. Dozier, J. 2006. Conceptual management plan for the Tom Yawkey Wildlife Center. South Carolina Department of Natural Resources, Columbia, SC. 151 pp. Dufrêne, M., and P. Legendre. 1997. Species assemblages and indicator species: The need for a flexible asymmetrical approach. Ecological Monographs 67:3 45–366. Southeastern Naturalist 291 C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 Dumond, M., and M. Villard. 2001. Does Coyote diet vary seasonally between a protected and an unprotected forest landscape? Ecoscience 8:301–310. Eskew, T.S. 2012. Best-management practices for reducing Coyote depredation on Loggerhead Sea Turtles in South Carolina. Clemson University, Clemson, SC. 105 pp. Farias, A.A., and M.J. Kittlein. 2008. Small-scale spatial variability in the diet of Pampas Foxes (Pseudalopex gymnocercus) and human-induced changes in prey base. Ecological Research 23:543–550. Gipson, P.S. 1974. Food habits of Coyotes in Arkansas. Journal of Wildlife Management 38:848–853. Godbois, I.A., L.M. Conner, B.D. Leopold, and R.J. Warren. 2005. Effect of diet on mass loss of Bobcat scat after exposure to field conditions. Wildlife Society Bulletin 33:149–153. Grigione, M.M., P. Burman, S. Clavio, S.J. Harper, D. Manning, and R.J. Sarno. 2011. Diet of Florida Coyotes in a protected wildland and suburban habitat. Urban Ecosystems 14:655–663. Hall, D. 1979. An ecological study of the Coyote-like canid in Louisiana. M.Sc. Thesis. Louisiana State University, Baton Rouge, LA. 233 pp. Hilderbrand, G.V., S. Farley, C. Robbins, T. Hanley, K. Titus, and C. Servheen. 1996. Use of stable isotopes to determine diets of living and extinct bears. Canadian Journal of Zoology 74:2080–2088. Hinton, J.W., M.J. Chamberlain, and F.T. van Manen. 2012. Long-distance movements of transient Coyotes in Eastern North Carolina. American Midland Naturalist 168:281–288. Hoerath, J.D. 1990. Influences of Coyotes on game animals as monitored by fecal analysis. M.Sc. Thesis. Auburn University, Auburn, AL. Hopkins-Murphy, S. 1989. The Santee Delta-Cape Romain Unit of the Carolinian-South Atlantic Biosphere Reserve. Pp. 79–91, In W.P. Gregg, S.L. Krugman, and J.D. Wood (Eds.). Proceedings of the Symposium on Biosphere Reserves. US Department of the Interior, National Park Service, Atlanta, GA. 291 pp. Kilgo, J.C., H.S. Ray, M. Vukovich, M.J. Goode, and C. Ruth. 2012. Predation by Coyotes on White-tailed Deer neonates in South Carolina. The Journal of Wildlife Management 76:1420–1430. Lander, M.E., T.R. Loughlin, M.G. Logsdon, G.R. VanBlaricom, B.S. Fadely, and L.W. Fritz. 2009. Regional differences in the spatial and temporal heterogeneity of oceanographic habitat used by Steller Sea Lions. Ecological Applications 19:1645–1659. Lavin, S.R., T.R. Van Deelen, P.W. Brown, R.E. Warner, and S.H. Ambrose. 2003. Prey use by Red Foxes (Vulpes vulpes) in urban and rural areas of Illinois. Canadian Journal of Zoology 81:1070–1082. Lea, M., C. Guinet, Y. Cherel, M. Hindell, L. Dubroca, and S. Thalmann. 2008. Colonybased foraging segregation by Antarctic Fur Seals at the Kerguelen Archipelago. Marine Ecology-Progress Series 358:273–287. Litvaitis, J.A., and J.H. Shaw. 1980. Coyote movements, habitat use, and food habits in southwestern Oklahoma. Journal of Wildlife Management 44:62–68. Livingston, T.R., P.S. Gipson, W.B. Ballard, D.M. Sanchez, and P.R. Krausman. 2005. Scat removal: A source of bias in feces-related studies. Wildlife Society Bulletin 33:172–178. Martin, A.C., and W.D. Barkley. 1961. Seed Identification Manual. University of California Press, Berkley, CA. 221 pp. Marucco, F., D.H. Pletscher, and L. Boitani. 2008. Accuracy of scat sampling for carnivore diet analysis: Wolves in the Alps as a case study. Journal of Mammalogy 89:665–673. Southeastern Naturalist C.R. Etheredge, S.E. Wiggers, O.E. Souther, L. Lagman, G. Yarrow, and J. Dozier 2015 Vol. 14, No. 2 292 McCune, B., J.B. Grace, and D.L. Urban. 2002. Analysis of Ecological Communities. Volume 28. MjM Software Design, Gleneden Beach, OR. 300 pp. Michaelson, K. 1975. Food habits of Coyotes in northwestern Louisiana. M.Sc. Thesis. Louisiana Tech University, Ruston, LA. 28 pp. Mielke, P.W., Jr., and K.J. Berry. 2001. Permutation Methods: A Distance-function Approach. Springer Series in Statistics, New York, NY. 344 pp. Miller, J.H., and K.V. Miller. 2005. Forest Plants of the Southeast and Their Wildlife Uses. University of Georgia Press, Athens, GA. 454 pp. Moore, T.D., L.E. Spence, and C.E. Dugnolle. 1974. Identification of the dorsal guard hairs of some mammals of Wyoming. Wyoming Game and Fish Department Bulletin Number 14. 177 pp. Morey, P.S., E.M. Gese, and S. Gehrt. 2007. Spatial and temporal variation in the diet of Coyotes in the Chicago metropolitan area. The American Midland Naturalist 158:147–161. Murie, O.J., and M. Elbroch. 2005. A Field Guide to Animal Tracks. Volume 3. Houghton Mifflin, New York, NY. 391 pp. Parker, G. 1995. Eastern Coyote: The Story of its Success. Nimbus Publishing, Halifax, NS, Canada. 254 pp. Roest, A.I. 1986. Key-guide to Mammal Skulls and Lower Jaws. Mad River Press, Inc., Eureka, CA. 39 pp. Rühe, F., M. Ksinsik, and C. Kiffner. 2008. Conversion factors in carnivore-scat analysis: Sources of bias. Wildlife Biology 14:500–506. Sanchez, D.M., P.R. Krausman, T.R. Livingston, and P.S. Gipson. 2004. Persistence of carnivore scat in the Sonoran Desert. Wildlife Society Bulletin 32:366–372. Schrecengost, J.D., J.C. Kilgo, D. Mallard, H.S. Ray, and K.V. Miller. 2008. Seasonal food habits of the Coyote in the South Carolina coastal plain. Southeastern Naturalist 7:135–144. Scott, S.D., and C. McFarland. 2010. Bird Feathers: A Guide to North American Species. Stackpole Books, Mechanicsburg, PA. 358 pp. Smith, R.A., and M.L. Kennedy. 1983. Food habits of the Coyote (Canis latrans) in western Tennessee. Journal of the Tennessee Academy of Science 58:27–30. Wagner, G.D., and E.P. Hill. 1994. Evaluation of southeastern Coyote diets during the Wild Turkey reproductive season. Proceedings of the Annual Conference of Southeastern Association of Fish and Wildlife Agencies 48:173–181. Wooding, J.B., E.P. Hill, and P.W. Sumner. 1984. Coyote food habits in Mississippi and Alabama. Proceedings of the Annual Conference of Southeastern Association of Fish and Wildlife Agencies 38:182–188. Yarrow, G.K., and D. Yarrow. 1999. Managing Wildlife. Sweetwater Press, Birmingham, AL. 588 pp.