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Prey Selection by Three Mesopredators that are Thought to Prey on Eastern Wild Turkeys (Meleagris gallopavo sylvestris) in the Pineywoods of East Texas
Haemish I.A.S. Melville, Warren C.Conway, Michael L.Morrison, Christopher E. Comer, and Jason B. Hardin

Southeastern Naturalist, Volume 14, Issue 3 (2015): 447–472

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Southeastern Naturalist 447 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 22001155 SOUTHEASTERN NATURALIST 1V4o(3l.) :1444,7 N–4o7. 23 Prey Selection by Three Mesopredators that are Thought to Prey on Eastern Wild Turkeys (Meleagris gallopavo sylvestris) in the Pineywoods of East Texas Haemish I.A.S. Melville1,2,*, Warren C.Conway3, Michael L.Morrison1, Christopher E. Comer4, and Jason B. Hardin5 Abstract - Predation, especially during the nesting and poult-rearing seasons, may inhibit Meleagris gallopavo (Wild Turkey) recruitment in east Texas. Numerous authors cite Lynx rufus (Bobcat), Canis latrans (Coyote), and Procyon lotor (Raccoon) as predators of Wild Turkey. Consequently, we investigated prey selection of these 3 common mesopredators using scat analysis. We also investigated prey-population dynamics using capture–mark– recapture techniques for small mammals (Rodentia), and spotlight surveys and track plate counts for Sylvilagus floridanus (Eastern Cottontail). We found no evidence that mesopredators preyed upon Wild Turkeys. Small mammals and lagomorphs were the primary components of mesopredator diets. Small-mammal numbers varied seasonally; however, Cottontail relative abundance did not. Mesopredator diets were most diverse in summer. In summer, Bobcats increased their use of small mammals, whereas Coyotes and Raccoons diversified their diets to include seasonal fruits. Decline in small-mammal populations and increase in mesopredator dietary diversity coincided with Wild Turkey nesting and poultrearing seasons, which potentially could result in an increased threat to Wild Turkeys during the nesting and poult-rearing season. Introduction Meleagris gallopavo L. (Wild Turkey) used to occupy approximately 12 million hectares in East Texas (Campo and Dickson 1990). Overharvesting and habitat destruction led to their local extinction by 1883 (Campo and Dickson 1990, Newman 1945). Restoration attempts in this region have been ongoing since 1924 (Newman 1945). Although most attempts to reestablish populations of Wild Turkeys in the United States have been successful, this has not been the case in east Texas (Boyd and Oglesby 1975, Isabelle 2010, Lopez et al. 2000, Newman 1945) where predation by mesopredators including Lynx rufus Schreber (Bobcat), Canis latrans Say (Coyote), and Procyon lotor L. (Raccoon) are thought to confound reintroduction programs of Wild Turkeys (George 1997, Kelly 2001). Predation is considered the primary cause of mortality for Wild Turkeys (Hamilton and Vangilder 1992, Hughes et al. 2005, Kennamer 2005, Miller and Leopold 1Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station, TX 77843. 2Current address - No. 8 25th Street, Zakher, Al Ain, United Arab Emirates. 3Department of Natural Resources Management, Texas Tech University, MS 2125, Lubbock, TX 79409-2125. 4Arthur Temple College of Forestry and Agriculture, Stephen F. Austin State University, Nacogdoches, TX 75962. 5Texas Parks and Wildlife Department, Oakwood, TX 75855. *Corresponding author - haemish.melville@gmail.com. Manuscript Editor: Renn Tumlison Southeastern Naturalist H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 448 1992). Female Meleagris gallopavo sylvestris Vieillot (Eastern Wild Turkeys), their eggs, and poults are particularly susceptible to predation during nesting and brood rearing (Lopez et al. 2000, Speake 1980). Predation may limit recruitment potential of low-density populations (Kot et al. 1993, Messier and Crête 1985, Newsome et al. 1989, Terborgh et al. 2001, Trout and Tittensor 1989), such as Wild Turkeys in East Texas (Isabelle 2010). Coyotes, Bobcats, and Raccoons live sympatrically over large portions of their range, including east Texas. Bobcats are carnivorous (Anderson 1985, Anderson and Lavallo 2003), whereas Coyotes and Raccoons are omnivorous (Schmidly 2004). Small mammals (Bartoszewicz et al. 2008, Fedriani et al. 2000, Litvaitis and Harrison 1989) and lagomorphs (Azevedo et al. 2006, Bekoff and Gese 2003, Gehrt 2003, Schmidly 2004) are primary components of the diets of these mesopredators. Behavioral mechanisms, including diet selection and use of space facilitate co-existence among mesopredators (Wang and Macdonald 2009). Predators adapt their feeding habits to the availability of suitable prey (Krebs 1978, Sunquist and Sunquist 1989). Several mechanisms affect mesopredator prey selection. Predators display functional (Baker et al. 2001, Boutin 1995, Holling 1959) as well as behavioral and numerical responses (Dunn 1977; Jędrzejewska and Jędrzejewski 1998; Schmidt and Ostfeld 2003, 2008) to changes in prey availability. Temporal variation in resource availability is a feature of the natural environment (Holt 2008). Smallmammal populations vary seasonally (O’Connell 1989) and annually (Windberg 1998). It seems that mesopredators respond to variations in prey availability, consuming preferred prey when they are available and diversifying their diets when preferred prey are scarce (Schoener 1971) . To gauge the effect of predation by mesopredators on Eastern Wild Turkeys in the Pineywoods of east Texas, we investigated the following hypotheses: (1) Eastern Wild Turkeys contribute significantly to the diets of Bobcats, Coyotes, and Raccoons; (2) mesopredator diets vary seasonally; (3) mesopredator diets overlap significantly; (4) mesopredator prey availability varies seasonally; (5) predators respond functionally to changes in prey availability; and (6) low points of prey populations coincide with the Wild Turkey nesting season. Our results will contribute to the understanding of the threat posed by these mesopredators to Eastern Wild Turkeys in east Texas. Study area We conducted our study on 2 sites from January 2009 to September 2011 in the Pineywoods Ecoregion of Nacogdoches and Angelina counties, east Texas. Study site 1 (31°30'N, 94°43'W) encompassed 1360 ha and was intensively managed for wildlife using controlled burns on a 5-year rotational basis. Study site 2 (31°25'N, 94°23'W) encompassed 5000 ha and was managed for timber, with fire excluded from the system. We chose these study sites because they were areas where stable Wild Turkey populations existed and they represented major land uses in east Texas. Southeastern Naturalist 449 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 The Pineywoods occur in the western Gulf coastal region and extend through east Texas, northwestern Louisiana, and southwestern Arkansas. Much of the natural vegetation of the Pineywoods has been altered due to planting of Pinus taeda L. (Loblolly Pine) and the exclusion of fire (Omernik et al. 2008). Private land holdings account for 63% of the Pineywoods, much of which is in small parcels (0.4 to 3.6 ha). A consequence of this pattern of ownership is an increase in forest fragmentation (US Department of Agriculture 2002). Pineywoods forests that are not under silviculture typically consist of an overstory that includes pine and oak species in association with Liquidambar styraciflua L. (Sweet Gum), Carya spp. (hickories), Fraxinus americana L. (White Ash), and Ilex opaca Ait. (American Holly) and an understory including Ilex vomitoria Ait. (Yaupon), Cornus florida L. (Flowering Dogwood), Asimina triloba (L.) Dunal (Pawpaw), Callicarpa americana L. (American Beautyberry), Chasmanthium laxum var. sessiliflorum (L.) Yates (Longleaf Uniola), Rubus plicatus Weihe and Nees (Bramble Blackberry), and grasses including Panicum spp. Mean annual rainfall in the Pineywoods is 1192 mm, and monthly mean precipitation varies from a minimum of 55 mm in July to a maximum of 116.4 mm in May. The mean annual minimum and maximum temperatures are 12.8 °C and 25.5 °C, respectively, with a mean summer maximum of 35 °C (Sivanpillai et al. 2005). Methods We used the natural (delineated by solstices and equinoxes) seasons for all analyses (winter: 21 December to 20 March, spring: 21 March to 20 June, summer: 21 June to 20 September, fall: 21 September to 20 December) because this schedule is relevant to the habits of mesopredators and prey species, including Eastern Wild Turkeys. We surveyed from winter 2009 (January) to summer 201 1 (August). Mesopredator dietary analysis To assess mesopredator dietary composition, we analyzed scat (Putman 1984) collected from Coyotes, Bobcats, and Raccoons. Scats collected from these species can be distinguished by appearance (Toweill and Anthony 1988). We searched several drainage lines and creek beds once every 2 weeks, and collected scats opportunistically on roads within study sites for the duration of the study. Scat collection was non-random with respect to microhabitat (Neale and Sacks 2001). After collection, we washed the scats and separated the contents into 4 categories: (1) bones and teeth, (2) hair and feathers, (3) plant material, and (4) insects. We identified scat-sample contents using osteological and hair keys, samples from the study sites, and reference books (Toweill and Anthony 1988). In addition, the microscopic characteristics of hairs were used to identify prey species (Prugh 2005). We made cuticular-scale impressions on microscope slides (Melville et al. 2004), examined them under compound microscopes, and compared them to keys and reference slides to identify the taxa represented by the hair. Hair medullary shapes from scat samples were compared to a key (Debelica and Thies 2009), and tooth and bone remains to reference material. When identificaton of Southeastern Naturalist H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 450 samples could not be determined using standard references, we consulted specialist taxonomists. Prey population monitoring Small mammals. Live-trapping grids (100 traps, 100 m x 100 m) and capture– mark–recapture (CMR) methodologies were used to estimate small-mammal populations (Edalgo and Anderson 2007, Parmenter et al. 2003, Reed et al. 2007, Wiewel et al. 2007). Annually, we randomly selected 6 of a total of 29 one-yearold Eastern Wild Turkey nest locations (3 on each study site) for grid placement. Nests were located during a study investigating the nesting ecology of Eastern Wild Turkeys in the Pineywoods of east Texas that ran concurrently with ours. We also selected 6 random locations (3 on each study site) for a total of 12 CMR surveys every season; 24 survey grids for the study. We sampled the same grids seasonally and changed grid locations annually to ensure that the Wild Turkey nest sites were a maximum of 1 year old. A mixture of rolled oats and peanut butter was used as bait, and each grid was sampled for 7 consecutive nights. Each animal that was captured was implanted with a uniquely numbered passive integrated transponder (PIT) tag (Biomark®). Our small-mammal capture records were used to estimate population size of the 3 most abundant small-mammal species: Reithrodontomys fulvescens J.A. Allen (Fulvous Harvest Mouse), Sigmodon hispidus Say and Ord (Hispid Cotton Rat), and Peromyscus leucopus Rafinesque (White-footed Mouse), for each trapping grid using the robust design method in program MARK (White and Burnham 1999). To ensure independence of samples, we positioned trapping grids in year 2 at least 1 km from any previous grid locations, and the grids on nest sites were associated with new Wild Turkey nests. Lagomorphs. We used spotlight counts and track-plate surveys to monitor seasonal variation in abundance of Sylvilagus floridanus J.A. Allen (Eastern Cottontail) (Malaney and Frey 2006, Williams et al. 2012). The 2 complimentary monitoring protocols were implemented seasonally because track plates have not been validated to monitor changes in relative abundance of lagomorphs (Ray and Zielinski 2008). A spotlight route was established on each study site. Routes traversed the length and breadth of each study site. Each spotlight count was conducted by 2 observers, each equipped with a spotlight and were initiated immediately after sunset, counting all observable Cottontails. Each spotlight route was approximately 20 km long (19 km on study site 2, 22 km on study site 1). We drove each route on alternate evenings for 14 days (7 iterations per study site) each season and conducted consecutive spotlight counts on individual study sites travelling in the opposite direction to the previous count. Each route was driven at a constant speed of 10 km/h. When bad weather curtailed counts, we delayed the spotlight surveys until the next evening (Fletcher et al. 1999). We conducted track-plate surveys comprising 25 track plates laid out in 2 parallel lines of 12 and 13 on each site. We positioned track plates along the same Southeastern Naturalist 451 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 roads as those used during spotlight counts to facilitate direct comparison between methods. Each track plate was located at least 150 m from the road. Track plates were constructed by covering 1 side of plywood backing plates (0.5 m x 0.5 m) with a Biofoam® impression surface (Hooper and Rea 2009). We placed track plates at least 320 m apart (i.e., greater than 1 Eastern Cottontail home-range diameter; Bond et al. 2002) so that each track plate could be considered an independent sample (Hamm et al. 2003, Ray and Zielinski 2008). Track plates were deployed simultaneously on both sites and exposed for 14 consecutive days. We checked the track plates every alternate day (7 times during each exposure period) and recorded any fresh track impressions. Fresh tracks, irrespective of whether they comprised a single impression or several tracks, were recorded as a single detection for each species on each track plate (Ray and Zielinski 2008, Sar geant et al. 1998). Analyses Mesopredator diet. We estimated relative frequency of each prey item within mesopredator scats by species, study site, year, and season. We used chi-square analysis (Zar 1999) to estimate differences among categories of dietary components among and within species, between years, between sites, and among seasons (Fedriani et al. 2000). We calculated a Shannon-Wiener index of dietary diversity for each mesopredator species (Jethva and Jhala 2004). Dietary overlap was calculated using Pianka’s index (O; Fedriani et al. 2000, Glen and Dickman 2008, Pianka 1973) for each species pair (Bobcat and Coyote, Bobcat and Raccoon, Coyote and Raccoon); we considered overlap index values >0.6 to be biologically significant (Bethea et al. 2006, Pianka 1976, Wallace 1981). Prey populations Small mammals. We calculated minimum known alive (Krebs 1966, Merritt et al. 2001) values for each species on each grid seasonally, and used factorial analysis of variance (ANOVA; Zar 1999) to compare differences between years, seasons, study sites, species, and site types (nest site versus random site). We utilized Pollock’s Robust Design (Pollock 1982) in program MARK (White and Burnham 1999) to estimate populations of the 3 most commonly captured species: Fulvous Harvest Mice, Hispid Cotton Rats, and White-footed Mice. Primary sampling intervals were set at 7 consecutive days per season; secondary samples were individual days within a primary period. We used these population estimates to compare species by year, study site, season, and nest site versus random site using factorial ANOVA (Zar 1999). Lagomorphs. We used spotlight count data to calculate a spotlight count index, and the number of Eastern Cottontail detections (R)/km to index relative abundance. These index values were used to compare lagomorph abundance between years, study sites, and seasons with factorial ANOVA (Zar 1999). We calculated a track-plate index by dividing the number of track-plate detections (RT) per unit time by number of track plates (TP) (Crooks 2002, Dijak and Thompson 2000, Drennan et al. 1998, Glennon et al. 2002, Lenth et al. 2008, Southeastern Naturalist H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 452 Wilson and Delahay 2001) on a site and seasonal basis. Index values between year, season, and study site (Gentry and Vierling 2007) were compared using factorial ANOVA (Zar 1999). We utilized a logistic regression model including the variables year, study site, and season relative to detection of Eastern Cottontails on track plates and validated it with Hosmer-Lemeshow and likelihood ratio tests. Linear regression was used to determine if spotlight and track-plate indices correlated (Schmidt et al. 2011). We evaluated all statistical tests at α = 0.05. Results Mesopredator diets We collected and analyzed 1764 mesopredator scats from January 2009 to August 2011: Bobcat, n = 637 (fall = 27, spring = 160, summer = 152, winter = 298); Coyote, n = 841 (fall = 54, spring = 235, summer = 252, winter = 299); and Raccoon, n = 286 (fall = 28, spring = 123, summer = 71, winter = 64). During this study 3383 prey occurrences (Bobcat = 976, Coyote = 1688, Raccoon = 719) were recorded (Melville 2012). Bobcats. Forty types of ingesta were identified, comprising vegetable matter, insects, fishes, birds, reptiles, and mammals (Appendix 1). Despite finding avian remains in Bobcat scats, no Wild Turkey remains were found in these samples. Bobcat diets did not differ between study sites (χ2 = 2.89, df = 21, P = 0.175). Bobcat diets did not vary between 2010 and 2011 (χ2 = 16.21, df = 13, P = 0.238), but did vary between 2009 and 2010 (χ2 = 22.92, df = 13, P = 0.043), effected largely by differences in consumption of Odocoileus virginianus Zimmerman (White-tailed Deer; 2009 = 10%, 2010 = 7.8%), lagomorphs (2009 = 24.3%, 2010 = 37.7%), and small mammals (2009 = 49%, 2010 = 37.5%) including Hispid Cotton Rats (2009 = 28.4%, 2010 = 19.6%). Between 2009 and 2011, major differences in consumption (χ2 = 47.98, df = 14, P < 0.001) were effected by insects (2009 = 1.4%, 2011 5.6%), White-tailed Deer (2009 = 10%, 2011 = 6.7%), lagomorphs (2009 = 24.3%, 2011 = 45.4%), and small mammals (2009 = 49%, 2011 = 27.7%) including Hispid Cotton Rats (2009 = 28.4%, 2011 = 20.2%). Annual Bobcat diets did not differ from fall or spring diets; however, they differed from summer and winter diets (Table 1). Bobcat diets varied seasonally, with the greatest contribution of White-tailed Deer (16.4%) in summer. Lagomorphs were the primary food source for Bobcats, with the greatest contribution in the fall (41.1%). Hispid Cotton Rats were consistently present in Bobcat diets, but contributed most (32.4%) in the winter. Some seasonal fruits were detected in Bobcat diets—notably Vitis rotundifolia Michx. (Muscadine Grape; 3%) in summer (Appendix 1). Spring and fall diets were similar (Table 1). Diversity values for Bobcat diets varied (fall H = 1.93, summer H = 2.47; Table 2). Coyotes. Forty-nine types of ingesta, including vegetable matter, insects, fishes, birds, reptiles, and human-made goods (Appendix 2) were present in Coyote scats. There were no Eastern Wild Turkey remains in Coyote scats, despite finding other avian prey remains in samples. Coyote diets differed between sites (χ2 = 43.97, df = 14, P < 0.001) and years (2009 vs 2010: χ2 = 75.37, df = 17, P < 0.001; 2009 Southeastern Naturalist 453 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 vs 2011: χ2 = 120.77, df = 17, P < 0.001; 2010 vs 2011: χ2 = 119.79, df = 17, P < 0.001). The dietary components that contributed most to the annual variation were lagomorphs (2009 = 13.1%, 2010 = 12.4%, 2011 = 22.4%), and small mammals (2009 = 19.5%, 2010 = 13.0%, 2011 = 7.7%), in particular Hispid Cotton Rats (2009 = 11.8%; 2010 = 8.1%; 2011 = 5.1%). Annual diets differed from seasonal diets, and seasonal diets differed (Table 1). Various dietary components peaked Table 1. Chi-square test results comparing the seasonal diets of 3 mesocarnivores in the Pineywoods of east Texas from January 2009 to August 2011. Species Seasons compared χ2 df P Bobcat Annual vs fall 19.44 14 0.149 Bobcat Annual vs spring 19.69 14 0.140 Bobcat Annual vs summer 39.51 14 <0.001 Bobcat Annual vs winter 29.30 14 0.009 Bobcat Fall vs spring 15.17 14 0.367 Bobcat Fall vs summer 27.18 12 0.007 Bobcat Fall vs winter 32.92 10 <0.001 Bobcat Spring vs summer 39.55 14 <0.001 Bobcat Spring vs winter 50.25 14 <0.001 Bobcat Summer vs winter 68.82 12 <0.001 Coyote Annual vs fall 68.79 19 <0.001 Coyote Annual vs spring 205.29 15 <0.001 Coyote Annual vs summer 165.66 19 <0.001 Coyote Annual vs winter 223.84 16 <0.001 Coyote Fall vs spring 211.73 15 <0.001 Coyote Fall vs summer 72.80 18 <0.001 Coyote Fall vs winter 152.12 13 <0.001 Coyote Spring vs summer 342.26 15 <0.001 Coyote Spring vs winter 260.34 13 <0.001 Coyote Summer vs winter 322.28 14 <0.001 Raccoon Annual vs fall 73.88 13 <0.001 Raccoon Annual vs spring 66.16 13 <0.001 Raccoon Annual vs summer 31.18 19 <0.001 Raccoon Annual vs winter 68.97 15 <0.001 Raccoon Fall vs spring 167.34 13 <0.001 Raccoon Fall vs summer 45.83 16 <0.001 Raccoon Fall vs winter 83.43 13 <0.001 Raccoon Spring vs summer 97.84 17 <0.001 Raccoon Spring vs winter 120.73 16 <0.001 Raccoon Summer vs winter 106.31 20 <0.001 Table 2. Shannon Wiener diversity index (H) values for the diets of 3 mesocarnivores in the Pineywoods of east Texas from January 2009 to August 2011. Season Bobcat (H) Coyote (H) Raccoon (H) Annual 2.35 2.71 2.75 Fall 1.93 2.44 2.43 Spring 2.29 2.12 1.96 Summer 2.47 2.90 2.83 Winter 2.13 2.09 2.31 Southeastern Naturalist H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 454 seasonally in Coyote diets: White-tailed Deer (28.5%), Sus scrofa L.(Feral Hog; 18%), and Hispid Cotton Rats (12.6%) in winter, and lagomorphs (19.7%) in spring (Appendix 2). Diversity values for Coyote diets varied seasonally (winter H = 2.08, summer H = 2.90; Table 2). Raccoons. Forty-one types of ingesta including vegetable matter, insects, fishes, birds, mammals, reptiles, aquatic invertebrates, and human-made items (Appendix 3) were present in Raccoon scats. No Eastern Wild Turkey remains were found in Raccoon scats, despite there being remnants of other avian prey in the samples. Raccoon diets did not differ between sites (χ2 = 19.03, df = 11, P = 0.061). Raccoon diets differed between years (2009 vs 2010: χ2 = 23.17, df = 12, P = 0.026; 2009 vs 2011: χ2 = 45.34, df = 12, P < 0.001; 2010 vs 2011: χ2 = 67.62, df = 12, P < 0.001). Dietary components that contributed most to the annual variation included insects (2009 = 14.3%, 2010 = 11%, 2011 = 35.3%), White-tailed Deer (2009 = 5.3%, 2010 = 12.7%, 2011 = 2.9%), lagomorphs (2009 = 7.3%, 2010 = 5%, 2011 = 15.4%), Hispid Cotton Rats (2009 = 7.3%, 2010 = 5.1%, 2011 = 2.9%), and Cambarus sp (Crawfish) (2009 = 2%, 2010 = 2.5%, 2011 = 5.9%). Annual diets differed from seasonal diets, and diets differed by season (Table 1). Various dietary components peaked seasonally: insects in spring (24.5%) and winter (23.7%), White-tailed Deer (12.7%) in winter, Hispid Cotton Rats in spring (7.5%) and summer (7%), and Crawfish (6.8%) in winter. Diversity values for Raccoon diets varied seasonally (spring H = 1.96, summer H = 2.83; Table 2). Comparison of mesopredators dietary composition. Diets of Bobcats, Coyotes, and Raccoons differed annually and seasonally (Table 3). Pianka’s overlap index values indicated that diets of mesopredators overlapped to varying degrees of biological significance (O > 0.6; Table 3). Bobcat diets overlapped greatly with Coyote diets annually (O = 0.72) and seasonally (spring [O = 0.65] and summer [O = 0.68]). Bobcat and Raccoon diets did not overlap. Coyote diets overlapped greatly with Raccoon diets annually (O = 0.69) and seasonally (spring [O = 0.88] and summer [O = 0.76]). Table 3. Chi-square test results and the associated Pianka dietary overlap (O) values for 3 mesocarnivores in the Pineywoods of east Texas from January 2009 to August 2011. Species Compared Season χ2 df P O Bobcat vs Coyote Annual 570.29 14 <0.001 0.72 Bobcat vs Coyote Fall 42.13 10 <0.001 0.58 Bobcat vs Coyote Spring 144.08 13 <0.001 0.65 Bobcat vs Coyote Summer 148.51 12 <0.001 0.68 Bobcat vs Coyote Winter 243.47 10 <0.001 0.58 Raccoon vs Coyote Annual 471.35 20 <0.001 0.69 Raccoon vs Coyote Fall 30.53 10 <0.001 0.30 Raccoon vs Coyote Spring 87.79 14 <0.001 0.88 Raccoon vs Coyote Summer 107.82 17 <0.001 0.76 Raccoon vs Coyote Winter 326.97 14 <0.001 0.40 Bobcat vs Raccoon Annual 746.07 20 <0.001 0.41 Bobcat vs Raccoon Fall 58.35 10 <0.001 0.19 Bobcat vs Raccoon Spring 206.39 14 <0.001 0.39 Bobcat vs Raccoon Summer 180.47 18 <0.001 0.45 Bobcat vs Raccoon Winter 344.24 11 <0.001 0.27 Southeastern Naturalist 455 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 Prey Small mammals. We captured 1922 small mammals (representing 7 species) during 67,200 trap nights from January 2009 to December 2010 (Table 4). Number of species captured varied between sample sites (F = 25.01; df = 5, 342; P < 0.001). We captured 562 White-footed Mice, 42 Blarina brevicauda Say (Short-tailed Shrew), 276 Hispid Cotton Rats, 863 Fulvous Harvest Mice, 76 Ochrotomys nuttalli Harlan (Golden Mouse), 12 Neotoma floridana Ord (Eastern Woodrat) and 90 Peromyscus gossypinus LeConte (Cotton Mouse). We detected seasonal differences in small-mammal captures (F = 16.37; df = 3, 344; P < 0.001; winter n = 855 vs summer n = 330, Tukey HSD = -1.045, P < 0.001; winter n = 855 vs fall n = 250, Tukey HSD = -1.393, P < 0.001; spring n = 678 vs summer n= 330, Tukey HSD = -0.701, P = 0.009; spring n = 678 vs fall n = 250, Tukey HSD = -1.05, P < 0.001). (Fig. 1). There was no variation in small-mammal captures between years (F = 0.117; df = 1, 346; P = 0.733), study sites (F = 0.108; df = 1, 346; P = 0.743), or grid types (nest or random) (F = 2.233; df = 1, 346; P = 0.136). Table 4. Number of small mammals captured during a capture–mark–recapture survey in the Pineywoods of east Texas, from January 2009 to December 2010. Common name Scientific name Total captures White-footed Mouse Peromyscus leucopus 563 Short-tailed Shrew Blarina carolinensis 42 Hispid Cotton Rat Sigmodon hispidus 276 Fulvous Harvest Mouse Reithrodontomys fulvescens 863 Golden Mouse Ochrotomys nuttalli 76 Eastern Woodrat Neotoma floridana 12 Cotton Mouse Peromyscus gossypinus 90 Total 1922 Figure 1. Trends in seasonal numbers of small mammals captured during a mark–recapture survey in the Pineywoods of east Texas, January 2009–December 2010. Southeastern Naturalist H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 456 Population estimates for the 3 most commonly captured species of small mammals did not vary between years (Fig. 2). Harvest Mouse populations were greatest during winter (mean = 20.5, SE = 2.86) and spring (mean = 19.1, SE = 2.91), then declined during the summer months (mean = 5.4, SE = 0.95) into fall (mean = 4.3, SE = 0.97) (Table 5, Fig. 2). Numbers of White-footed Mice varied between study sites (site 1 mean = 5.8, SE = 0.84 vs. study site 2 mean = 9.8, SE = 1.29). Whitefooted Mice followed similar seasonal patterns to Harvest Mice, (winter mean = 14.3, SE = 1.83; spring mean = 7.76, SE = 1.37; summer mean = 4.0, SE = 0.73; and fall mean =4.9, SE = 1.15; Table 5; Fig. 2). Hispid Cotton Rat populations varied by grid type, with greater numbers occurring on random sites (mean = 7.6, SE = 1.36) than on Wild Turkey nest sites (mean = 3.2, SE = 0.47). Numbers of Hispid Cotton Rats varied seasonally (summer mean = 9.7, SE = 2.3; fall mean = 2.6, SE = 0.8; winter mean = 3.4, SE = 2.3; and spring mean = 4.4, SE = 1.0; Table 5, Fig. 2). Lagomorphs. We conducted 70 spotlight counts over 1433 km (study site 2: 769 km, study site 1: 664 km) seasonally from April 2010 to August 2011, and recorded 132 Eastern Cottontails (study site 2 = 100, study site 1 = 32; Table 6). We observed no difference in numbers of Eastern Cottontails between years (F = 0.025; df = 1, 68; P = 0.874) or seasons (F = 1.56; df = 3, 66; P = 0.207). Spotlight indexes differed between sites (study site 2: mean = 0.2 R/km, SE = 0.02; study site 1: mean = 0.04 R/km, SE = 0.01; F = 23.27; df = 1, 68; P < 0.001; Fig. 3). Figure 2. Mean population numbers (± 1 SE) for the 3 most abundant small mammals captured in the Pineywoods of east Texas, January 2009–December 2010. Southeastern Naturalist 457 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 We recorded 3500 track-plate nights for lagomorphs from April 2010 to August 2011, and detected Eastern Cottontails 121 times (study site 2: n = 73; study site 1: n = 48). We converted these to an index of detections per track plate (RT/ TP, range = 0.0–0.2; Table 7). There was no difference in track-plate indexes between years (F = 0.87; df = 1, 68; P = 0.354) or seasons (F = 1.63; df = 3, 66; P = 0.191), but we detected a difference between sites (study site 2: mean = 0.08 RT/ TP, SE = 0.01; study site 1: mean = 0.06 RT/TP, SE = 0.01; F = 4.371; df = 1, 68; P = 0.04; Fig. 4). We used logistic regression to model the influence of the variables year, study site, and season on detection of lagomorph on the track-plates. Detection of tracks was influenced by study site (Z = -2.344, df = 1, P = 0.091; Table 8). This finding Table 5. Results of ANOVA comparing the variables associated with the 3 most abundant species caught in the Pineywoods of east Texas from January 2009 to August 2011. Harvest Mouse Hispid Cotton Rat White-footed Mouse Comparisons F df P F df P F df P Study sites 0.757 1, 77 0.39 1.081 1, 41 0.31 6.201 1, 81 0.02 Years 0.436 1, 77 0.51 1.105 1, 41 0.30 0.308 1, 81 0.50 Grid types 0.351 1, 77 0.56 5.914 1, 41 0.02 0.107 1, 81 0.74 Seasons 18.340 3, 75 <0.001 4.476 3, 39 0.01 12.190 3, 79 <0.001 Tukey P Tukey P Tukey P Winter vs spring -0.274 0.92 -0.550 0.56 -0.953 0.02 Winter vs summer -2.241 <0.001 0.416 0.76 -1.733 <0.001 Winter vs fall -2.574 <0.001 -1.065 0.08 -1.599 <0.001 Spring vs summer -1.966 <0.001 0.968 0.12 -0.779 0.08 Spring vs fall -2.299 <0.001 -0.513 0.64 -0.646 0.20 Summer vs fall -0.333 0.87 -1.481 0.01 0.133 0.98 Table 6. Spotlight index values (rabbits per km) for detections of Sylvilagus floridanus (Eastern Cottontail Rabbit) in the Pineywoods of east Texas from spring 2010 to summer 2011. Site Season n SE Spotlight index Both All 70 0.01 0.10 Site 1 All 35 0.02 0.15 Site 2 All 35 0.01 0.04 Both Fall 14 0.02 0.05 Site 1 Fall 7 0.03 0.04 Site 2 Fall 7 0.03 0.06 Both Spring 28 0.02 0.10 Site 1 Spring 14 0.02 0.06 Site 2 Spring 14 0.03 0.15 Both Summer 14 0.05 0.13 Site 1 Summer 7 0.01 0.01 Site 2 Summer 7 0.06 0.26 Both Winter 14 0.02 0.87 Site 1 Winter 7 0.02 0.04 Site 2 Winter 7 0.04 0.14 Southeastern Naturalist H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 458 was confirmed by a likelihood ratio test (log likelihood = -434.05, df = 5, P = 0.037), and a Hosmer-Lemeshow goodness-of-fit test (χ2 = 6.89, df = 8, P = 0.55). There was no correlation between our spotlight and track-plate indices (F = 1.349; df = 1, 68; P = 0.25; r2 = 0.02). Discussion Bobcats, Coyotes, and Raccoons had diverse diets in the Pineywoods of east Texas. Their diets included a variety of mammals, insects, birds, plants, fishes, and reptiles, but no evidence of Eastern Wild Turkey remains. Our results support previous studies in the Southeast that have found limited support for the supposition Figure 3. Spotlight count index (rabbits/km) for 2 study sites in the Pineywoods of east Texas, April 2010–August 2011. Table 7. Track plate index (rabbit impressions per track plate) for detections of Sylvilagus floridanus (Eastern Cottontail Rabbit) in the Pineywoods of east Texas from spring 2010 to summer 2011. Site Season n SE Track index Both All 70 0.01 0.07 Site 1 All 35 0.01 0.08 Site 2 All 35 0.01 0.06 Both Fall 14 0.01 0.05 Site 1 Fall 7 0.01 0.05 Site 2 Fall 7 0.02 0.06 Both Spring 28 0.01 0.06 Site 1 Spring 14 0.02 0.05 Site 2 Spring 14 0.02 0.07 Both Summer 14 0.02 0.08 Site 1 Summer 7 0.02 0.04 Site 2 Summer 7 0.02 0.13 Both Winter 14 0.01 0.09 Site 1 Winter 7 0.02 0.08 Site 2 Winter 7 0.01 0.10 Southeastern Naturalist 459 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 that mesopredators prey on Wild Turkeys, even in areas with abundant populations of this potential prey (Chamberlain and Leopold 1999, Leopold and Chamberlain 2002, Wagner and Hill 1994). When mesopredators prey on large eggs, they seldom ingest entire egg shells (Larivière 1999). Even if mesopredators consumed large numbers of eggs, it would be unlikely that we would have found many eggshell fragments in scats. It is also unlikely that we would have detected poults in scats (Wagner and Hill 1994). Thus, despite the lack of evidence, it is possible that mesopredators preyed on Eastern Wild Turkey eggs and poults. Numerous investigations of Wild Turkey mortality indicate that predation is the most important cause of reduced survival (Isabelle 2010, Kurzejeski et al. 1987, Miller et al. 1998, Speake et al. 1985, Swank et al. 1985), yet there is conflicting evidence relating to the importance of predation by mesopredators on Wild Turkeys. Our results in east Texas confirm the findings of other studies that Wild Turkeys do not contribute greatly to the diets of mesopredators (Chamberlain and Leopold 1999, Leopold and Chamberlain 2002, Melville 2012, Wagner and Hill 1994). Figure 4. Eastern Cottontail Rabbit track index (tracks/plate/night) calculated for 2 study sites in the Pineywoods of east Texas, April 2010–August 2011. Table 8. Confidence intervals from logistic regression of the variables associated with likelihood of detecting Sylvilagus floridanus (Eastern Cottontail Rabbit) tracks on track plates in the Pineywoods of east Texas from Spring 2010 to Summer 2011. *** = highly significant; * = significant. Variable Lower Upper Z df P (Intercept) -2.838 -1.353 -5.54 1 <0.001 *** Site = Site 1 -0.834 -0.077 -2.34 1 0.019 * 2011 -0.629 0.629 0.00 1 1.000 Spring -1.042 0.109 -1.56 1 0.118 Summer -0.929 0.711 -0.26 1 0.795 Fall -1.496 0.241 -1.41 1 0.159 Southeastern Naturalist H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 460 Small mammals contributed greatly to the diets of mesopredators in the Pineywoods of east Texas, although their populations declined from spring into summer during 2009 and 2010. During spring and summer, mesopredators increased diversity in their diets. Increased dietary diversity has been linked to limitation in food availability (Clavero et al. 2003). Decline in prey availability occurred while Wild Turkeys were nesting and raising poults. The synchrony between the decline in prey populations and nesting of Wild Turkeys may have increased the probability of Wild Turkey nest and poult predation, especially considering that mesopredators diversified their diets during this period. Despite this, we found no Wild Turkey remains in mesopredator scats. Sympatric predators display variability in dietary breadth and overlap consistent with principles of resource partitioning (Azevedo et al. 2006). Our results for Bobcats, Coyotes, and Raccoons in the Pineywoods confirms previous findings that the degree of dietary overlap between mesopredators varies seasonally (Azevedo et al. 2006, Chamberlain and Leopold 1999, Fedriani et al. 2000, Major and Sherburne 1987). Levels of dietary overlap between Bobcats and Coyotes were marginally less than what is considered biologically significant (O > 0.6) in fall and winter, but overlap increased to biologically significant levels in spring and summer. Similar patterns of seasonal variation in dietary overlap between Bobcats and Coyotes have been observed in central Mississippi (Chamberlain and Leopold 1999) and California (Fedriani et al. 2000). An even more pronounced seasonal change in significance of overlap was evident between Coyotes and Raccoons. There was no biologically significant dietary overlap in winter or fall; however, dietary overlap was significant in spring and summer. This change in overlap was probably due to their selection of seasonal fruits. Few other studies have investigated dietary overlap between Coyotes and Raccoons. Where comparisons have been made, they were conducted on an annual basis and provided results that showed little overlap between Coyote and Raccoon diets (Azevedo et al. 2006). Although a seasonal trend of increasing dietary overlap was evident between Bobcats and Raccoons, the level of overlap was never biologically significant. The increase in dietary overlap between mesopredators in spring and summer, combined with high dietary diversity index values in summer may indicate seasonal resource limitation in terms of food availability during those times of year (Clavero et al. 2003). Recent weather events can affect small-mammal communities (French et al. 1976, Grant et al. 1985, Schmidt and Ostfeld 2008). Our study of small mammals spanned 2009 and 2010, a period associated with the onset of a 500-year drought in Texas from 2010–2011 (CIESS 2012). The mechanism of influence of harsh weather on small-mammal populations occurs indirectly through the impact on vegetation productivity (Grant et al. 1985). Despite this, we found no evidence of a general decline in small-mammal populations in 2010. Our results are similar to those of Grant et al. (1985) that found no effect of temperature and precipitation on Hispid Cotton Rat and White-footed Mouse populations. Grant et al. (1985) also found that Fulvous Harvest Mouse populations were influenced by temperature fluctuations, but not precipitation. In the Pineywoods, Fulvous Harvest Mouse numbers declined Southeastern Naturalist 461 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 consistently during the warmer months. It is unlikely that the full effect on small mammals of the Texas drought of 2010–2011 was fully identified during our study because this would only have been evident after our monitoring ceased. Bobcats, Coyotes, and Raccoons preyed on small mammals to varying degrees. The only small-mammal species that contributed substantially to mesopredator diets was the Hispid Cotton Rat, a primary prey item for Bobcats. Other small mammals were preyed upon opportunistically by mesopredators. Mesopredators are known to limit Hispid Cotton Rat populations, especially in areas where fire is excluded (Conner et al. 2011). This may be the case in the Pineywoods, especially on commercial timber sites where fire has been excluded as a management tool. It is unknown whether mesopredators limit other small mammals. Mesopredators seemed to respond to variation in prey availability. Bobcats responded to changes in Hispid Cotton Rat availability by increasing their consumption of them when the population increased, similar to what was observed in Georgia (Baker et al. 2001). Lagomorphs were present in the diets of the mesopredators under study. Coyotes and Bobcats used lagomorphs extensively, despite their apparent low densities. Cottontails seemed to be the most important dietary item for Bobcats, consistently comprising more than 20% of the items in their scats, although their frequency seemed to vary relative to availability of Hispid Cotton Rats. Coyotes and Raccoons are opportunistic, generalist predators, varying their food intake relative to availability (Bekoff and Gese 2003, Gehrt 2003). During spring and summer, Coyote and Raccoon diets contained a large percentage of fruit. Locating and consuming fruit probably required less energy than searching for and capturing live prey. Thus, Coyotes and Raccoons likely improved foraging efficiency by including a high proportion of fruit when seasonally abundant (Chamberlain and Leopold 1999, MacArthur and Pianka 1966, Norber g 1977). Eastern Wild Turkeys did not contribute to diets of Bobcats, Coyotes, or Raccoons in east Texas during our study. Our study did not prove that the predators under study did not use Eastern Wild Turkeys, but our data did not reveal use during our study. It seems that mesopredators responded to seasonal variations in resource availability, and that Eastern Cottontails, despite occurring in seemingly low densities, contributed greatly to their diets. We suggest that further studies be undertaken to determine if the mesopredators in question prey extensively on nests and poults of Eastern Wild Turkeys in the Pineywoods. Further, we suggest that more investigations into the population dynamics of Eastern Cottontails in east Texas are warranted. Acknowledgments Our project was funded by federal excise taxes on sport-hunting arms and ammunition, Grant W132R, with the Texas Parks and Wildlife Department, as well as by Texas A&M University (Department of Wildlife and Fisheries Science), Stephen F. 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Kelly. 2012. Monitoring translocated riparian Brush Rabbits and surveying for and censusing of Brush Rabbits and woodrats. Endangered Species Recovery Program, California State University, Stanislaus Turlock, CA. 23 pp. Wilson, G.J., and R.J. Delahay. 2001. A review of methods to estimate the abundance of terrestrial carnivores using field signs and observation. Wildlife Research 28:151–164. Windberg, L.A. 1998. Population trends and habitat associations of rodents in southern Texas. American Midland Naturalist 140:153–160. Zar, J.H. 1999. Biostatistical Analysis (Fourth Edition). Prentice-Hall, NJ. 663 pp. Southeastern Naturalist 467 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 Appendix 1. The occurrence of dietary items in the Lynx rufus (Bobcat) scats collected in the Pineywoods of east Texas, January 2009–August 2011. * indicates the presence of Bobcat hair in their own scats, which was attributed to grooming. Annual 2009 2010 2011 Fall Spring Summer Winter Type/species Common name n % n % n % n % n % n % n % n % Bird Cardinalis cardinalis Cardinal 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 Gallus gallus domesticus Chicken 24 2.5 16 2.5 6 2.9 2 1.7 4 10.3 5 2.0 7 3.0 8 1.8 Picoides sp. Woodpecker 1 0.1 0 0.0 1 0.5 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 Unidentified bird 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 1 0.2 Fish Unidentified fish 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 2 0.8 0 0.0 0 0.0 Insects Unidentified Insects 19 2.0 9 1.4 3 1.5 7 5.9 2 5.1 12 4.9 1 0.4 4 0.9 Mammals Odocoileus virginianus White tailed deer 88 9.1 64 10.0 16 7.8 8 6.7 3 7.7 26 10.6 38 16.4 21 4.7 Ovis aries Sheep 1 0.1 0 0.0 0 0.0 1 0.8 0 0.0 1 0.4 0 0.0 0 0.0 Sus scrofa Hog 12 1.2 8 1.3 3 1.5 1 0.8 0 0.0 4 1.6 2 0.9 6 1.3 Dasypus novemcinctus Armadillo 4 0.4 2 0.3 2 1.0 0 0.0 0 0.0 1 0.4 3 1.3 0 0.0 Lynx rufus Bobcat 9* 6* 2* 1* 0* 2* 2* 5* Felis sylvestris catus Domestic Cat 3 0.3 2 0.3 1 0.5 0 0.0 0 0.0 1 0.4 0 0.0 2 0.4 Urocyon cinereoargenteus Gray Fox 7 0.7 4 0.6 1 0.5 2 1.7 0 0.0 2 0.8 1 0.4 4 0.9 Didelphis virginiana Opossum 3 0.3 2 0.3 1 0.5 0 0.0 0 0.0 1 0.4 0 0.0 2 0.4 Procyon lotor Raccoon 29 3.0 19 3.0 5 2.5 5 4.2 0 0.0 9 3.7 11 4.7 9 2.0 Vulpes vulpes Red Fox 2 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 1 0.4 0 0.0 Sylvilagus floridanus Eastern Cottontail 245 25.3 136 21.3 58 28.4 51 42.9 15 38.5 73 29.8 42 18.1 115 25.5 Sylvilagus aquaticus Swamp Rabbit 41 4.2 19 3.0 19 9.3 3 2.5 1 2.6 13 5.3 5 2.2 22 4.9 Sciurus niger Eastern Fox Squirrel 3 0.3 2 0.3 1 0.5 0 0.0 0 0.0 0 0.0 1 0.4 2 0.4 Sciurus carolinensis Eastern Gray Squirrel 1 0.1 0 0.0 1 0.5 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 Neotoma floridana Eastern Woodrat 69 7.1 48 7.5 17 8.3 4 3.4 3 7.7 14 5.7 15 6.5 37 8.2 Sigmodon hispidus Hispid Cotton Rat 246 25.4 182 28.4 40 19.6 24 20.2 6 15.4 47 19.2 47 20.3 146 32.4 Rattus norvegicus Norway Rat 6 0.6 2 0.3 3 1.5 1 0.8 0 0.0 2 0.8 1 0.4 3 0.7 Rattus rattus Rat 2 0.2 0 0.0 0 0.0 0 0.0 1 2.6 0 0.0 0 0.0 1 0.2 Southeastern Naturalist H.I.A.S. 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Hardin 2015 Vol. 14, No. 3 468 Annual 2009 2010 2011 Fall Spring Summer Winter Type/species Common name n % n % n % n % n % n % n % n % Peromyscus gossypinus Cotton Mouse 20 2.1 16 2.5 4 2.0 0 0.0 0 0.0 4 1.6 7 3.0 9 2.0 Peromyscus leucopus White-footed Mouse 21 2.2 15 2.3 3 1.5 3 2.5 1 2.6 6 2.4 2 0.9 12 2.7 Ochrotomys nuttalli Golden Mouse 7 0.7 6 0.9 1 0.5 0 0.0 0 0.0 0 0.0 2 0.9 5 1.1 Reithrodontomys humulis Eastern Harvest Mouse 20 2.1 19 3.0 1 0.5 0 0.0 0 0.0 3 1.2 9 3.9 8 1.8 Reithrodontomys fulvescens Fulvous Harvest Mouse 34 3.5 26 4.1 7 3.4 1 0.8 1 2.6 4 1.6 10 4.3 19 4.2 Snake Coluber spp. Racer species 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 Nerodia spp. Watersnake species 1 0.1 1 0.2 0 0.0 0 0.0 1 2.6 0 0.0 0 0.0 0 0.0 Agkistrodon piscivorus Cotton Mouth 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0 Crotalus horridus Timber Rattlesnake 1 0.1 0 0.0 1 0.5 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 Plant Callicarpa americana Beauty Berry 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 Rubus plicatus Bramble Blackberry 2 0.2 1 0.2 0 0.0 1 0.8 0 0.0 2 0.8 0 0.0 0 0.0 Prunus virginiana Choke Cherry 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 0 0.0 2 0.9 0 0.0 Unidentified grass 35 3.6 21 3.3 9 4.4 5 4.2 1 2.6 11 4.5 11 4.7 12 2.7 Smilax rotundifolia Greenbriar 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 0 0.0 2 0.9 0 0.0 Vitis rotundifolia Muscadine 7 0.7 7 1.1 0 0.0 0 0.0 0 0.0 0 0.0 7 3.0 0 0.0 Pyrus spp. Pear 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 Southeastern Naturalist 469 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 Appendix 2. The occurrence of dietary items in the Canis latrans (Coyote) scats collected in the Pineywood of east Texas, January 2009–August 2011. * indicates the presence of Coyote hair in their own scats, which was attributed to grooming. Annual 2009 2010 2011 Fall Spring Summer Winter Type/species Common name n % n % n % n % n % n % n % n % Bird Cardinalis cardinalis Cardinal 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 Gallus gallus domesticus Chicken 41 2.5 15 1.5 19 4.8 7 2.6 5 5.2 3 0.6 12 2.1 21 4.4 Zenaida macroura Mourning Dove 2 0.1 1 0.1 0 0.0 1 0.4 0 0.0 0 0.0 0 0.0 2 0.4 Unidentified bird 10 0.6 5 0.5 4 1.0 1 0.4 2 2.1 4 0.8 4 0.7 0 0.0 Fish Unidentified fish 2 0.1 2 0.2 0 0.0 0 0.0 0 0.0 1 0.2 1 0.2 0 0.0 Insect Unidentified Insect 70 4.2 25 2.5 10 2.5 35 12.9 0 0.0 55 10.6 11 2.0 4 0.8 Mammal Odocoileus virginianus White-tailed Deer 292 17.7 170 17.2 70 17.8 52 19.1 22 22.7 79 15.3 55 9.8 136 28.5 Equus caballus Horse 1 0.1 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0 0 0.0 0 0.0 Sus scrofa Hog 145 8.8 92 9.3 26 6.6 27 9.9 6 6.2 26 5.0 27 4.8 86 18.0 Ovis aries Sheep 1 0.1 0 0.0 0 0.0 1 0.4 0 0.0 1 0.2 0 0.0 0 0.0 Lynx rufus Bobcat 5 0.3 3 0.3 2 0.5 0 0.0 0 0.0 1 0.2 4 0.7 0 0.0 Canis latrans Coyote 34* 21* 9* 4* 1* 13* 9* 11* Urocyon cinereoargenteus Gray Fox 9 0.5 5 0.5 3 0.8 1 0.4 0 0.0 1 0.2 4 0.7 4 0.8 Didelphis virginiana Opossum 2 0.1 1 0.1 0 0.0 1 0.4 0 0.0 0 0.0 0 0.0 1 0.2 Procyon lotor Raccoon 38 2.3 16 1.6 12 3.1 10 3.7 1 1.0 9 1.7 15 2.7 13 2.7 Dasypus novemcinctus Armadillo 21 1.3 7 0.7 12 3.1 2 0.7 0 0.0 7 1.4 13 2.3 1 0.2 Lepus californicus Black-tailed Jack Rabbit 2 0.1 1 0.1 1 0.3 0 0.0 0 0.0 1 0.2 0 0.0 1 0.2 Sylvilagus floridanus Eastern Cottontail 218 13.2 120 12.2 43 10.9 55 20.2 11 11.3 94 18.2 43 7.7 70 14.7 Sylvilagus aquaticus Swamp Rabbit 21 1.3 9 0.9 6 1.5 6 2.2 0 0.0 8 1.5 7 1.2 6 1.3 Neotoma floridana Eastern Woodrat 33 2.0 21 2.1 9 2.3 3 1.1 1 1.0 10 1.9 8 1.4 14 2.9 Rattus norvegicus Norway Rat 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 Sigmodon hispidus Hispid Cotton Rat 162 9.8 116 11.8 32 8.1 14 5.1 11 11.3 42 8.1 49 8.7 60 12.6 Peromyscus gossypinus Cotton Mouse 11 0.7 9 0.9 1 0.3 1 0.4 1 1.0 2 0.4 5 0.9 3 0.6 Peromyscus leucopus White-footed Mouse 9 0.5 6 0.6 3 0.8 0 0.0 0 0.0 2 0.4 3 0.5 4 0.8 Southeastern Naturalist H.I.A.S. 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Hardin 2015 Vol. 14, No. 3 470 Annual 2009 2010 2011 Fall Spring Summer Winter Type/species Common name n % n % n % n % n % n % n % n % Ochrotomys nuttalli Golden Mouse 5 0.3 3 0.3 2 0.5 0 0.0 0 0.0 0 0.0 4 0.7 1 0.2 Reithrodontomys humulis Eastern Harvest Mouse 14 0.8 13 1.3 0 0.0 1 0.4 0 0.0 2 0.4 8 1.4 4 0.8 Reithrodontomys fulvescens Fulvous Harvest Mouse 27 1.6 21 2.1 4 1.0 2 0.7 3 3.1 3 0.6 10 1.8 11 2.3 Mus muscullus House Mouse 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0 Geomys breviceps Baird’s Pocket Gopher 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0 Plant Callicarpa americana Beauty Berry 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0 Rubus plicatus Bramble Blackberry 174 10.6 114 11.6 21 5.3 39 14.3 0 0.0 141 27.3 33 5.9 0 0.0 Prunus virginiana Choke Cherry 37 2.2 26 2.6 11 2.8 0 0.0 3 3.1 0 0.0 34 6.1 0 0.0 Zea mays Corn 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0 Unidentified grasses 90 5.5 63 6.4 19 4.8 8 2.9 3 3.1 20 3.9 42 7.5 25 5.2 Smilax rotundifolia Green briar 58 3.5 40 4.1 17 4.3 1 0.4 7 7.2 1 0.2 49 8.7 1 0.2 Morus nigra Mulberry 9 0.5 5 0.5 4 1.0 0 0.0 6 6.2 0 0.0 3 0.5 0 0.0 Vitis rotundifolia Muscadine 86 5.2 54 5.5 31 7.9 1 0.4 5 5.2 0 0.0 81 14.4 5 1.0 Pyrus spp. Pear 1 0.1 0 0.0 1 0.3 0 0.0 1 1.0 0 0.0 0 0.0 0 0.0 Ampelopsis arborea Peppervine 5 0.3 0 0.0 5 1.3 0 0.0 0 0.0 0 0.0 5 0.9 0 0.0 Diospyros virginiana Persimone 21 1.3 11 1.1 10 2.5 0 0.0 8 8.2 0 0.0 13 2.3 0 0.0 Unidentified plants 8 0.5 0 0.0 8 2.0 0 0.0 0 0.0 0 0.0 8 1.4 0 0.0 Ligustrum vulgare Privet 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0 Cucurbita spp. Pumpkin 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0 Citrullus lanatus Watermelon 4 0.2 1 0.1 2 0.5 1 0.4 0 0.0 0 0.0 4 0.7 0 0.0 Reptile Unidentified lizard 1 0.1 0 0.0 1 0.3 0 0.0 1 1.0 0 0.0 0 0.0 0 0.0 Unidentified snakes 2 0.1 1 0.1 0 0.0 1 0.4 0 0.0 2 0.4 0 0.0 0 0.0 Miscellaneous Leather 2 0.1 2 0.2 0 0.0 0 0.0 0 0.0 2 0.4 0 0.0 0 0.0 Latex 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 Plastic 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 Southeastern Naturalist 471 H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin 2015 Vol. 14, No. 3 Appendix 3. The occurrence of dietary items in the Procyon lotor (Raccoon) scats collected in the Pineywood of east Texas, January 2009–August 2011. * indicates the presence of Raccoon hair in their own scats, which was attributed to grooming. Annual 2009 2010 2011 Fall Spring Summer Winter Type/species Common name n % n % n % n % n % n % n % n % Bird Gallus gallus domesticus Chicken 8 1.4 5 1.7 2 1.7 1 0.7 1 1.8 1 0.4 5 3.5 1 0.8 Picoides sp. Woodpecker 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 Unidentified bird 5 0.9 2 0.7 1 0.8 2 1.5 0 0.0 2 0.8 1 0.7 2 1.7 Fish Unidentified fish 4 0.7 3 1.0 1 0.8 0 0.0 0 0.0 1 0.4 3 2.1 0 0.0 Insect Unidentified insect 104 18.7 43 14.3 13 11.0 48 35.3 1 1.8 59 24.5 16 11.2 28 23.7 Mammal Odocoileus virginianus White-tailed Deer 35 6.3 16 5.3 15 12.7 4 2.9 5 8.9 11 4.6 7 4.9 15 12.7 Sus scrofa Feral Hog 9 1.6 5 1.7 2 1.7 2 1.5 0 0.0 4 1.7 2 1.4 3 2.5 Dasypus novemcinctus Armadillo 11 2.0 6 2.0 4 3.4 1 0.7 0 0.0 8 3.3 3 2.1 0 0.0 Procyon lotor Raccoon 162* 84* 42* 36* 11* 90* 45* 16* Urocyon cinereoargenteus Gray Fox 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 Sylvilagus floridanus EasternCottontail 47 8.5 21 7.0 5 4.2 21 15.4 2 3.6 24 10.0 9 6.3 12 10.2 Sylvilagus aquaticus Swamp Rabbit 2 0.4 1 0.3 1 0.8 0 0.0 0 0.0 0 0.0 2 1.4 0 0.0 Neotoma floridana Eastern Woodrat 4 0.7 1 0.3 3 2.5 0 0.0 1 1.8 2 0.8 1 0.7 0 0.0 Rattus norvegicus Norway Rat 1 0.2 0 0.0 1 0.8 0 0.0 1 1.8 0 0.0 0 0.0 0 0.0 Sigmodon hispidus Hispid Cotton Rat 32 5.8 22 7.3 6 5.1 4 2.9 1 1.8 18 7.5 10 7.0 3 2.5 Reithrodontomys fulvescens Fulvous Harvest Mouse 3 0.5 1 0.3 2 1.7 0 0.0 1 1.8 1 0.4 1 0.7 0 0.0 Ochrotomys nuttalli Golden Mouse 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0 Plant Callicarpa americana Beauty Berry 24 4.3 23 7.7 0 0.0 1 0.7 13 23.2 0 0.0 9 6.3 2 1.7 Rubus plicatus Bramble Blackberry 100 18.0 57 19.0 21 17.8 22 16.2 1 1.8 77 32.0 22 15.4 0 0.0 Celtis spp. 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 0 0.0 0 0.0 1 0.8 Prunus virginiana Choke Cherry 12 2.2 5 1.7 7 5.9 0 0.0 6 10.7 1 0.4 5 3.5 0 0.0 Zea mays Corn 28 5.0 19 6.3 1 0.8 8 5.9 0 0.0 2 0.8 2 1.4 24 20.3 Unidnetified grasses 30 5.4 12 4.0 12 10.2 6 4.4 3 5.4 15 6.2 9 6.3 3 2.5 Southeastern Naturalist H.I.A.S. 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Hardin 2015 Vol. 14, No. 3 472 Annual 2009 2010 2011 Fall Spring Summer Winter Type/species Common name n % n % n % n % n % n % n % n % Smilax rotundifolia Greenbriar 15 2.7 15 5.0 0 0.0 0 0.0 6 10.7 0 0.0 9 6.3 0 0.0 Celtis occidentalis Hackberry 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 0 0.0 0 0.0 1 0.8 Ilex spp. Holly 3 0.5 2 0.7 0 0.0 1 0.7 0 0.0 0 0.0 0 0.0 3 2.5 Morus nigra Mulberry 4 0.7 3 1.0 0 0.0 1 0.7 2 3.6 1 0.4 0 0.0 1 0.8 Vitis rotundifolia Muscadine 15 2.7 8 2.7 7 5.9 0 0.0 2 3.6 0 0.0 13 9.1 0 0.0 Ampelopsis arborea Peppervine 2 0.4 1 0.3 1 0.8 0 0.0 1 1.8 0 0.0 1 0.7 0 0.0 Diospyros virginiana Persimmon 10 1.8 6 2.0 4 3.4 0 0.0 6 10.7 0 0.0 4 2.8 0 0.0 Ilex decidua Possum Hawe 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 1 0.4 0 0.0 0 0.0 Cucurbita spp. Pumpkin 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8 Lolium multiflorum Ryegrass 5 0.9 3 1.0 0 0.0 2 1.5 0 0.0 0 0.0 0 0.0 5 4.2 Citrullus lanatus Watermelon 2 0.4 0 0.0 2 1.7 0 0.0 0 0.0 0 0.0 2 1.4 0 0.0 Nyssa aquatica Water Tupelo 4 0.7 4 1.3 0 0.0 0 0.0 0 0.0 0 0.0 2 1.4 2 1.7 Unidentified plants 8 1.4 4 1.3 4 3.4 0 0.0 2 3.6 2 0.8 4 2.8 0 0.0 Invertebrates Millipede 1 0.2 0 0.0 0 0.0 0 0.0 1 1.8 0 0.0 0 0.0 0 0.0 Cambarus spp. Crawfish 17 3.1 6 2.0 3 2.5 8 5.9 0 0.0 8 3.3 1 0.7 8 6.8 Aquatic snail 2 0.4 2 0.7 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 2 1.7 Micellaneous Duck tape 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0 Styrofoam 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 1