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An Assessment of Herpetofaunal and Non-Volant Mammal Communities at Sites in the Piedmont of North Carolina
Joshua M. Kapfer and David J. Muñoz

Southeastern Naturalist, Volume 11, Issue 1 (2012): 65–88

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2012 SOUTHEASTERN NATURALIST 11(1):65–88 An Assessment of Herpetofaunal and Non-Volant Mammal Communities at Sites in the Piedmont of North Carolina Joshua M. Kapfer1,2,* and David J. Muñoz1 Abstract - The southeastern United States contains a rich diversity of vertebrate species. Despite this, the Piedmont province of the southeastern US has received less attention than the more biologically diverse Coastal Plain and Mountain regions. Yet, the Piedmont region experiences the greatest anthropogenic impact and should be the focus of conservation efforts. In an attempt to obtain diversity information for this under-studied region, we surveyed amphibian, reptile, and non-volant mammal communities for one year at two sites in the Piedmont of North Carolina. Our survey methodologies included drift fences, artificial cover objects, camera traps, and visual encounter surveys. We captured or obtained evidence of a total of 49 species across both sites (mammals = 20, amphibians = 15, reptiles = 14), and over 2000 animals were captured or detected. We calculated measures of species richness, abundance, diversity, and evenness for each study site, and calculated similarity between sites. Diversity and evenness measures varied, but were generally highest for amphibians or reptiles and lowest for mammals. Measures of similarity between study sites indicated high similarity. The species we observed were comparable to those reported by past inventory projects in the Piedmont of North Carolina, although such projects have been sparse. Our results provide much-needed information on vertebrate communities in this under-studied region. Introduction The southeastern United States contains a rich diversity of native vertebrate species. North Carolina, for example, is home to over 200 species of amphibians, reptiles, and terrestrial mammals (Beane et al. 2010, Webster et al. 1985). Unfortunately, urbanization as a result of an expanding human population occurs at an alarming rate in North Carolina. Recent data regarding human population growth in the United States from 2008 to 2009 revealed that North Carolina was one of the fastest growing states (US Census Bureau 2010a). Furthermore, data collected from 2000 to 2010 suggests that the human population in North Carolina has increased by 18.5% (compared to 9.7% for the entire USA). It is also a densely populated state, with estimates of 195.8 people per square mi in 2010, compared to 87.3 people per square mi for the entire United States (US Census Bureau 2010b). Such high population densities and rapid growth results in a concomitant loss of habitat, as natural landscapes are converted to urban and suburban environments to meet the needs of this growing population. Habitat fragmentation and destruction from conversion of natural areas to urban/suburban environments is one of the main reasons for the loss of biodiversity that 1Departments of Environmental Studies and Biology, Elon University, Elon, NC 27244. 2Current address - Department of Biological Sciences, University of Wisconsin-Whitewater, Whitewater, WI 53190. *Corresponding author - 66 Southeastern Naturalist Vol. 11, No. 1 occurs globally (McKinney 2006). As a result, current species-extinction rates are much higher than normal (Barnosky et al. 2011). The southern Piedmont province is a region of the southeastern United States that is situated between the Atlantic Coastal Plain and the Appalachian Mountains. It is a plateau that stretches from Virginia to Alabama and includes North Carolina (Fig. 1). Mild climate, relatively flat topography, and an abundance of water have resulted in rapid growth of industry and human populations in the region since the 19th century (reviewed by Conroy et al. 2003). As a result, much of the natural landscape once available for wildlife in the southern Piedmont has been lost. In particular, a substantial portion of primary forest has been cleared since the time of European settlement (Conroy et al. 2003). Unfortunately, fragmentation of forested landscapes will likely increase over time due to continued human population growth and associated urban/suburban sprawl in the Piedmont (Wear and Greis 2001). It is estimated that this region will be subject to the greatest loss of forested land among all regions of the Southern United States (Wear and Greis 2001). There is a need to document species diversity in the southern Piedmont given the rate of habitat loss in the region. Such information can provide baseline data on local communities should future extripations or extinctions occur. Thus, assessing plant and animal communities and identifying potentially rare species in locations that have not been investigated are important endeavors. Biological inventories, measures of species richness, and measures of biological diversity are crucial first steps in assessing the status and trends of wildlife communities, the ecological robustness of a given area, and the effective planning of conservation strategies (Dorcas et al. 2006, Kremen 1994, Primack 2010, Tuberville et al. 2005). Much published research exists on the herpetofaunal and mammal communities of the Mountains (e.g., Ford et al. 2000, Hicks and Pearson 2003, Kaminski et al. 2007) and Coastal Plain (e.g., Hutchens and DePerno 2009, Meyers and Pike 2006, Mitchell et al. 1995, Tuberville et al. 2005) provinces of the southeastern USA. On the other hand, comparatively little work has been conducted in the southern Piedmont. Existing literature includes some research projects focused on the ecology or natural history of various vertebrate groups within the southern Piedmont (e.g., Matthews 1990, Todd et al. 2003, Willson and Dorcas 2004), while some past projects have also attempted to compile inventory lists for locations within this region (Kalcounis-Rueppel et al. 2007a, Rice et al. 2001). A handful of studies have inventoried mammal and herpetofaunal species along the border between the Mountains and Piedmont in South Carolina (Dorcas et al. 2006, Dorcas et al. 2010, Webster 2005). Several others have attempted to compare vertebrate communities among habitat types (e.g., Atkeson and Johnson 1979, Metts et al. 2001), investigate the effects of logging and forest removal (e.g., Pagels et al. 1992), or examine the influence of anthropogenic disturbance on specific taxonomic groups in the Piedmont (Kalcounis-Rueppell 2007b, Price et al. 2006, Price et al. 2010). 2012 J.M. Kapfer and D.J. Muñoz 67 To our knowledge, few published studies have attempted to quantify communities of several broad vertebrate taxonomic groups (i.e., amphibians, reptiles, and mammals) at locations within the southern Piedmont. In an effort to add information on the wildlife communities in this region, we surveyed herpetofaunal and non-volant mammal communities at study sites in Alamance County, NC (Fig. 1). These results will help further document the species and vertebrate community diversity present in the under-studied southern Piedmont. Study Sites Surveys were conducted simultaneously at two locations in Alamance County, NC: a larger, less-disturbed site and a smaller, more-disturbed site (Fig. 1). The study location we characterized as large and less-disturbed was a roughly 80.82- ha natural area, which we refer to as the “Haw Site”. Historically the site was used as a grist mill and homestead throughout the 19th century, and was subject to agricultural activity over a portion of its area roughly 40 to 60 years ago (Ryan Kirk and David Vandermast, Elon University, Elon, NC, pers. comm.). Since this time, however, it has remained undisturbed, with only periodic mowing of a small area for hay. The study location we characterized as small and more disturbed was a 9.83-ha parcel owned by Elon University, which we refer to as the “Elon Site”. This site was also subject to agriculture in the same time period as the larger site (Ryan Kirk and David Vandermast, pers. comm.). Although it Figure 1. General location of study sites within the Piedmont Province of North Carolina (Alamance County). 68 Southeastern Naturalist Vol. 11, No. 1 has partially responded since this agricultural disturbance, it may be classified as exurban, and is currently more vulnerable to encroachment of anthropogenic development. A portion of this site has also been periodically mowed for hay. These sites exist approximately 4 km (straight-line distance) from each other, and the immediate landscapes they are associated with historically shared many similar habitat and topographical features. Methods Habitat assessment Available habitat present within the boundaries of both study locations was first assessed through visual surveys on-site during Fall 2009. The extent of available habitat at both locations was then assessed via aerial photograph interpretation in a geographical information system (GIS; ArcMap 9.3, ESRI, Redlands, CA) with subsequent ground-truthing. We analyzed the landscape within a 50-m buffer associated with the property boundary of each study site in an effort to assess land-use immediately adjacent to the monitored properties. Although this buffer distance was somewhat arbitrary, we believed it would encompass the majority of potential bouts of movement for most of the monitored species on-site (with the exception of very mobile medium and large mammals). Thus, based on (1) the presence of similar habitat types at each site (albeit in differing quantities) and (2) their close geographic relationship to each other, the probability that these sites historically shared many species in common was likely high. Geographical information system analyses and on-site assessment of the property revealed that the habitats present within each site included upland deciduous forest, lowland deciduous forest, riparian, grassland/old field, edge between woodland and grassland, and disturbed. Lowland deciduous forests were comprised of both mesic forest and alluvial forest vegetative species, whereas upland deciduous forests were dominated by oak-hickory communities (Spira 2011). Riparian habitat (terrestrial habitat within 10–15 m of stream banks) was almost exclusively wooded, with vegetative species similar to those in lowland deciduous forests. Both sites contained transmission line right-of-ways, which experienced moderate to low vehicular traffic as part of ongoing maintenance. Although old field/grassland habitats in the Piedmont exist due to past disturbance, this habitat at our study sites was not currently influenced by anthropogenic activities (aside from annual mowing). Therefore, we did not consider this type of habitat to be “disturbed” at the same level as areas of manicured lawns. GIS analyses revealed that the proportion of habitat types associated with each site varied. Upland deciduous hardwood forest was the dominant habitat at both sites (Figs. 2, 3). Disturbed and oldfield/grassland habitats were in greater proportion at the Elon Site than at the Haw Site (Figs. 2, 3). The proportion of lowland deciduous hardwood forest waas slightly larger at the Haw Site than at the Elon Site (Figs. 2, 3). We considered the three 2012 J.M. Kapfer and D.J. Muñoz 69 primary habitats on-site to be upland deciduous hardwood forest, lowland deciduous hardwood forest, and oldfield/grassland. Survey techniques A variety of survey techniques targeting amphibians, reptiles, and nonvolant mammals were implemented to increase the likelihood of effectively capturing or observing all species present within each study location (Ryan et al. 2002). As suggested by Osbourne et al. (2005), we also surveyed a variety of habitat types to most accurately sample the wildlife communities present onsite. Therefore, specific permanent survey methods (i.e., drift fences, camera traps, and artificial cover objects) were employed within each of the primary Figure 2. The amount of each habitat type present and specific survey equipment locations at the Elon Site (Alamance County, NC). 70 Southeastern Naturalist Vol. 11, No. 1 habitat types at both sites: upland deciduous hardwood forest, lowland deciduous hardwood forest, and grassland. Mobile survey methods (i.e., visual encounter surveys) were conducted throughout the site, regardless of habitat type. All captured animals that could be handled were photographed, weighed to the nearest 0.1 g, measured to the nearest 0.1 cm, and released. We did not mark captured individuals for future identification. Passive infrared (PIR) triggered camera traps are an effective methodology for monitoring medium and large mammals (reviewed by O’Connell et al. 2011). For the purpose of this study, camera traps (Bushnell Trophy Cam, Bushnell Company, Overland Park, KS) provided a non-invasive means by which mammals that are otherwise difficult to capture or observe could be surveyed. All camera traps possessed infrared flash (as opposed to traditional white-flash), which we believed would reduce the likelihood of camera avoidance by wary species (Cutler and Swann 1999). Camera traps were set to take pictures in “bursts” of three at every trigger event in order to increase the Figure 3. The amount of each habitat type present and specific survey equipment locations at the Haw Site (Alamance County, NC). 2012 J.M. Kapfer and D.J. Muñoz 71 chances of obtaining suitable photographs. Three camera traps (one in each primary habitat type on-site) were deployed in July 2010 at each study site. The general location for camera deployment within a broad habitat type was selected randomly; however, cameras were specifically placed in a location that exhibited high animal activity (i.e., wildlife trails, etc.) as suggested by other camera-trap studies (e.g., Wilson et al. 1996). Cameras were mounted on tree trunks in protective metal boxes close to the ground (height of 30–60 cm) to ensure that small and large species were equally likely to be captured (Kelly 2008). Because many medium and large mammals are active year round, camera- trap surveys were conducted continuously over an entire year (from July 2010 to July 2011). This extended duration also helped eliminate any potential seasonal effects that may influence the presence/absence of species that are active year round. Cameras were serviced weekly to exchange memory cards, check battery life, and assess potential theft attempts. We did not attempt to identify individual animals captured by camera traps. Thus, animals captured were counted as individual “passes” and summed by species. We used drift fences with associated live traps to survey for amphibians, reptiles, and small mammals (Heyer et al. 1994, Schemnitz 2005, Wilson et al. 1996). Drift fences were comprised of erosion-control material, and were installed in an “X” configuration with each arm pointing in a cardinal direction. Each arm of the fence was approximately 15 m, totaling 60 m of fence at each drift-fence location. Fences were dug at least 20 cm into the ground to ensure animals could not burrow underneath. Locations for fence construction were selected mostly at random within the three primary habitats, although consideration was given to the feasibility of installation (i.e., whether fences could physically fit within the area selected; Figs. 2, 3). Multiple trap types were deployed along drift fences to achieve the greatest success of capturing a variety of species (Todd et al. 2007). Pitfall traps (i.e., 5-gallon plastic buckets) were dug in so that the upper lip of the bucket was flush with the surface of the ground. A single pitfall trap per fence was placed where the four arms of the drift fence intersected. Wire funnel traps (i.e., minnow traps; Memphis Net and Twine, Memphis, TN) were placed on each side of the fence along the north and south sections, equaling four funnel traps per drift-fence location (Schemnitz 2005). Specially constructed wooden box traps were placed at the ends of the west and east sections of each fence, two box traps per drift-fence location. Box-trap dimensions were 60 cm x 60 cm x 90 cm, and designed to catch amphibians, reptiles, and small mammals, with specific focus on larger snake species. We conducted drift-fence surveys from late July/early August 2010 until early November 2010. At that time, capture rates dropped to zero, or near zero, for at least two weeks, and non-camera surveys were discontinued for the winter. Driftfence surveys were then re-instated in March 2011, which also yielded very few captures for several weeks. Therefore, we believe we did not miss any period of substantial amphibian, reptile, or small mammal activity during the time when drift-fence surveys had been discontinued. In 2011, drift fence surveys were 72 Southeastern Naturalist Vol. 11, No. 1 conducted until mid-July. Drift-fence traps were engaged once per week when animal activity was moderate (March–April, August–November), and 2–3 d per week (checked at 24-hr intervals) when activity peaked (May–July). Accidental trap mortality was low, but salvageable specimens were deposited at the North Carolina Museum of Natural Sciences (small mammals) or the University of Wisconsin-Whitewater (amphibians and reptiles). The use of artificial cover objects (ACOs) is a well-accepted passive survey methodology for studying herpetofauna (Heyer et al. 1994); we also observed that ACOs attracted many small-mammal species. Although Sherman traps were originally deployed for small mammals (July–November 2010), we found them to be a less effective tool for capturing small mammals than ACOs and drift fences, particularly given the effort required to bait and set Sherman traps. Therefore, we discontinued Sherman trap use after fall of 2010, and no Sherman trap results were included in our analyses. We deployed ACOs constructed of 60-cm x 90-cm x 0.63-cm sheets of plywood along transects within the three primary habitats on-site. The starting location for each transect was chosen randomly within selected habitats, and from this random point, 10 ACOs, each spaced 15 m apart, were laid along the transect line (Figs. 2, 3). We conducted standardized ACO surveys weekly, on a schedule that conformed with our drift-fence survey schedule. Visual-encounter surveys (VES) are an effective technique often employed to monitor biological communities (Heyer et al. 1994, Karns 1986). They are also an important technique to include when surveying for species that are unlikely to be captured via other survey techniques. We considered a variety of observations obtained during VES as acceptable for our dataset. These included: live animals; bones, antlers, or carcasses; and other signs of wildlife (i.e., tracks and scat). Evidence such as bones, carcasses, and animal signs, was removed or identified so that it was not counted in subsequent surveys. Only tracks and scat that could be positively identified were included in our analyses. Visual encounter surveys were conducted year round, although the amount of time spent on-site decreased during the winter when only camera traps were being serviced. All VES were conducted haphazardly, as time allowed, and while surveyors were walking between permanent survey stations (i.e., drift fences, ACOs, and camera traps). We standardized these surveys by person-hour (the number of hours spent surveying a site times the total number of people surveying). Measures of community diversity To assess the diversity of the focal vertebrate communities, we calculated several standard measures. Species richness (S; Krebs 1998) was determined by counting all species identified during surveys. We also tallied abundances, or the total number of individuals observed, for each species detected. To make our results comparable with a wider range of past and future studies, we calculated two indices of species diversity: Shannon-Weiner (H') and Brillouin’s (H) (Krebs 2012 J.M. Kapfer and D.J. Muñoz 73 1998). We selected these measures of diversity because they are sensitive to the inclusion of rare species, or species in low abundance. Diversity indices and associated confidence limits (90%) were obtained by bootstrapping the data 5000 times (Krebs 1998). We calculated a Smith and Wilson’s measure of species evenness (J') for taxa at both sites. This particular evenness estimate was selected because it is not influenced by high or low measures of S, while being equally sensitive to rare and common species (Krebs 1998). Based on a recent re-evaluation of species evenness measures, we also include Pielou’s measure (J) for taxa at both sites (Jost 2010). The similarity between the vertebrate communities of interest at each site was calculated using both the community percentage similarity index and Morista’s measure of similarity (Krebs 1998). All diversity measures were conducted separately for amphibians, reptiles, mammals, and for all taxonomic groups combined. We used the software Ecological Methodology V 7.1 (Exeter Software, Setauket, NY) to calculate diversity measures, with the exception of Pielou’s measure of evenness, which was calculated by hand from our Shannon-Weiner diversity indices. We generally summarized in which habitat each species was most often observed (Table 1). Although this is not a measure of habitat preference (i.e., habitat use vs. habitat availability), it gives a rough estimate of the habitats that each species was often associated with on-site. We also calculated capture probabilities for all taxa surveyed via the Royle and Nichols (2003) model designed for capture-only data in the program PRESENCE V. 3.1 (Hines 2006). Capture probabilities of medium and large mammals were calculated from daily presence-absence data obtained by camera traps, and we did not include data from surveys of tracks and sign in this analysis. Capture probabilities of amphibians, reptiles, and small mammals were calculated based on presenceabsence data combined for all survey methods used (visual encounters, drift fences, ACOs) per week. Results Survey effort varied slightly among techniques and sites, but was mostly consistent. All drift fences at the Elon Site were checked on a total of 44 occasions, while all drift fences at the Haw Site were checked on 37 occasions. We checked all ACOs on a total of 41 (Elon Site) to 47 (Haw Site) occasions. Camera traps were operational from 368 d (Elon Site) to 380 d (Haw Site). The number of person hours spent conducting visual surveys varied, primarily due to the difference in the sizes of each study site. The much larger Haw Site required more time to reach survey equipment, resulting in 316.5 total person hours spent conducting VES, while only 231.83 person hours were totaled at the smaller Elon Site. Effectiveness differed among survey methods. Infrared-triggered camera traps yielded the highest number of individual animal captures at each site, which is reasonable because they were deployed for a longer time (24 h/day, 74 Southeastern Naturalist Vol. 11, No. 1 year round) . However, they are only useful when surveying for medium and large mammals. Regarding small vertebrate surveys, some techniques were more effective than others. At the Haw Site, a total of 85 individual animals, representing 23 species, were captured via drift fences. Of these, more than ten captures were obtained for only one species, Carphophis amoenus Say (Eastern Worm Snake). A total of 92 individual animals, representing 9 species, were captured under ACOs at the Haw Site. Although the raw abundances were similar between these methods at the Haw Site, the majority of ACO captures (79% of captures) were of only two species: Peromyscus leucopus Rafinesque (White-footed Mouse; n = 27) and Plethodon cylindraceus Harlan (White-spotted Slimy Salamander; n = 46). At the Elon Site, drift fences captured 76 individuals across only 19 species, and ACOs resulted in the capture of only 31 individual animals across eight species. Furthermore, the majority of these ACO captures at the Elon Site (n = 21, or 67%) were of Eastern Worm Snakes. Eleven orders and suborders were represented at the end of our surveys, which included 49 species (mammals: n = 20; amphibians: n = 15; reptiles: n = 14) and 2118 individual animal observations or detections (Table 1, Appendix 1). A total of 34 species and 758 individual animals were observed at the smaller, more disturbed Elon Site (excluding Canis familiaris L. [Domestic Dog] and Felis catus L.[Domestic Cat]). Mammals comprised the majority of the species present at this site, and were in greatest overall abundance, followed by reptiles and amphibians, respectively (Table 1). These numbers were smaller than those obtained at the much larger Haw Site, which yielded a total of 41 species and 1357 individuals, with mammals again having the highest S and abundance (Table 1). In general, diversity indices calculated for the combination of all taxa (i.e., amphibians, reptiles, and mammals) across sites ranged from 2.22 to 2.55 (H), and 2.30 to 2.65 (H'). The highest calculated diversity index among taxa varied by site, but was either associated with amphibians or reptiles, while mammal diversity was always lowest (Table 1). Highest measures of evenness also varied between amphibians and reptiles by study site, and were always lowest for mammals (Table 1). Both of the employed measures of community similarity between the study sites indicated that resemblance was high in all taxa except amphibians (Table 1). Most amphibian species were found in association with forested habitat (upland and lowland) or wooded riparian areas. Reptile and mammal species were more equally spread among forest and grassland (Appendix 1). Despite the effort mounted, capture probabilities were small for most of the species identified (Appendix 1). Discussion Due to limited past work on herpetofaunal and mammal community diversity in the southern Piedmont, there are few studies to compare our results against. 2012 J.M. Kapfer and D.J. Muñoz 75 Table 1. Diversity measures (90% confidence limits, where applicable, derived from 5000 bootstrapped iterations) for amphibian, reptile, non-volant mammal, and overall sampled communities at two sites studied in the Piedmont of North Carolina (Alamance County). Elon Haw Mammals Amphibians Reptiles All taxa Mammals Amphibians Reptiles All taxa Species richness (S) 14 9 11 34 17 13 11 41 Abundance (n) 627 49 82 758 1139 148 70 1357 Shannon-Weiner index (H') 1.69 2.14 2.54 2.65 1.24 3.07 2.78 2.30 (1.56–1.82) (1.72–2.51) (2.26–2.79) (2.50–2.79) (1.38–1.34) (2.88–3.25) (2.50–3.02) (2.17–2.42) Brouillin’s index (H) 1.64 1.85 2.30 2.55 1.20 2.86 2.49 2.22 (1.52–1.76) (1.48–2.19) (2.06–2.53) (2.41–2.69) (1.10–1.30) (2.68–3.03) (2.24–2.70) (2.10–2.35) Smith and Wilson’s evenness (J) 0.186 0.546 0.429 0.277 0.223 0.607 0.528 0.338 Pielou’s evenness (J') 0.640 0.973 0.998 0.751 0.437 0.999 0.997 0.619 Elon/Haw Mammals Amphibians Reptiles All taxa Morista’s measure of similarity 0.98 0.23 0.96 0.97 Community % similarity index 82.35 31.06 72.66 75.41 76 Southeastern Naturalist Vol. 11, No. 1 Mammal species richness at our study sites was greater than mammal species richness detected at a nearby urban/suburban Piedmont site (S = 11, excluding feral cats; Kalcounis-Rueppell 2007a). Our species lists were similar to herpetofaunal inventory information from sites in the western Piedmont of North Carolina (Rice et al. 2001), with a few notable differences (see below). Our measures of H' were higher than those reported by Metts et al. (2001) for lowland amphibian and reptile species associated with streams and ponds in the southern Piedmont of South Carolina. This difference is understandable considering that we surveyed for both lowland and upland species, rather than only focusing on aquatic habitats. Calculated S values for our sites were substantially lower than those reported from three national parks in the Coastal Plain region of North Carolina (Tuberville et al. 2005). Our estimates of H' for amphibians and reptiles were similar to those reported by studies conducted in the Alligator River National Wildlife Refuge, located in the Coastal Plain region of North Carolina (Meyers and Pike 2006). Our measures of J', however, were generally lower than those reported in that same study. As Meyers and Pike (2006) state, the herpetofaunal diversity of this refuge is noticeably lower than the surrounding landscape. This fact would explain why our estimates of herpetofaunal diversity from a location in the Piedmont, which is generally considered to be less diverse than the Coastal Plain, are comparable to their results. Not only were our values of S greater at the Haw Site, but we also observed that the abundances of many species were much greater there than at the Elon Site. The calculated amphibian and reptile diversity indices were also higher for the Haw Site, which is larger and less-disturbed. The diversity indices calculated for mammals, as well as the combination of all taxa surveyed, were larger at the Elon Site. Given the substantial difference in the sizes of these two properties and the much higher proportion of disturbed habitat associated with the Elon Site, we were surprised that the communities surveyed were so similar (Morista’s measure of similarity = 0.97; community percent similarity index = 75.41). We will review several examples from our results that highlight interesting interactions within the communities we studied. Odocoileus virginianus Zimmermann (White-tailed Deer) and Procyon lotor L. (Raccoons) The number of White-tailed Deer observations was substantially higher at the Haw Site (Appendix 1). Roseberry and Woolf (1998) report that the amount of forest coverage at the landscape level influences White-tailed Deer population densities in a given area. They also mention that harvest by human hunters can play a substantial role in regulating deer densities. The Haw Site contains a much larger amount of recently undisturbed forested habitat than the Elon Site, providing extensive suitable habitat for White-tailed Deer (Figs. 2, 3). Evidence of past hunting activity by humans was found at both sites, but it likely had less impact on deer densities at the larger Haw Site. Although hunting is currently prohibited on both sites, there is greater human presence at the Haw Site (i.e., 2012 J.M. Kapfer and D.J. Muñoz 77 county employees and outdoor recreationalists) to deter would-be illegal hunters compared to the Elon Site. Due to the size of the Haw Site, deer that remain within park boundaries will also have a greater buffer from hunters on adjacent properties than at the smaller Elon Site. All of these factors, coupled with the lack of many potential predators at either site, may explain why White-tailed Deer were more often observed at the Haw Site than the Elon Site. In contrast to the White-tailed Deer, nearly double the observations of Raccoon were made at the Elon Site than the Haw Site. This species is subsidized by the activities of humans, and it can thrive in suburban and urban environments. In fact, several studies have reported high population densities of this species in areas associated with anthropogenic landscapes (reviewed by Hadidian et al. 2010). Although the Elon Site is not urban or suburban, it is best described as exurban, and is in much closer contact with developed land than the Haw Site. Considering the extent of land occupied by humans nearthe Elon Site (Fig. 2), we are not surprised by the high concentration of Raccoons there. Small-mammal species As with our study, past mammal inventories in the Piedmont of North Carolina have found that White-footed Mice are abundant (Kalcounis-Rueppell 2007a; Table 1). Given this species’ preference for woodlands (reviewed by Lackey et al. 1985), which were ample at our study sites, we are not surprised by their high abundance. Another woodland rodent species captured during our surveys, which is frequently sympatric with the White-footed Mouse, was Ochrotomys nuttalli Harlan (Golden Mouse). Unlike the White-footed Mouse, the Golden Mouse was captured infrequently during our surveys, and none were captured at the Elon Site. Pearson (1953) suggested an inverse relationship occurs between the densities of Golden Mice and White-footed Mice in areas of sympatry. Christopher and Barrett (2006) found that Golden Mice will increase their use of arboreal space when sympatric with dense populations of White-footed Mice, which may have reduced the likelihood of their capture in our drift fences and ACOs. Therefore, our infrequent Golden Mouse captures are reasonable given the large number of White-footed Mice that we detected. Low numbers of other species, such as Zapus hudsonius Zimmerman (Meadow Jumping Mouse), make sense given their preferences for open, grassy habitats (Whitaker 1972), which were not particularly abundant at our sites. This species, in particular, is seldom reported in the Piedmont region of North Carolina, which makes our observations valuable. For example, only 10 catalogued specimens of Meadow Jumping Mice from the Piedmont exist in the North Carolina Museum of Natural Sciences collection (NCSM 209–216, NCSM 423, and NCSM 475; Lisa Gatens, North Carolina Museum of Natural Sciences, Raleigh, NC, pers. comm.). Most of these specimens were collected from 40 to >100 years previously, and from locations several counties removed from our study sites. We are somewhat surprised by the sparseness of small-mammal captures at the smaller Elon Site, including White-footed Mice, and the absence of other species 78 Southeastern Naturalist Vol. 11, No. 1 often associated with disturbed landscapes (Appendix 1). These low numbers are in contrast to what past studies have suggested, which is that abundances of small mammals may be reduced in larger habitat fragments. This relationship occurs because these fragments are large enough for individuals to set up and defend territories, so a few dominant individuals exclude others (Foster and Gaines 1991). Winter mortality for the White-footed Mouse increases substantially in small woodland fragments (Wilder et al. 2005), which may have led to a lower population at the Elon Site. Furthermore, the relatively numerous observations of several important rodent predators (e.g., Canis latrans Say [Coyote; Bekoff and Gese 2003], Urocyon cinereoargenteus Schreber [Gray Fox; Fritzell and Haroldson 1982]) and the presence of Domestic Cats at the smaller Elon Site may have influenced rodent populations. Terrestrial salamander communities Terrestrial salamanders have been suggested as strong indicators of forest ecosystem health (Welsh and Droege 2001). In fact, Hicks and Pearson (2003) determined that terrestrial salamanders are even sensitive to historic alteration of woodland habitats. In general, the richness and abundance of salamanders was greater at the less disturbed Haw Site, which would indicate higher ecosystem health. Yet, it is interesting that Ambystoma opacum Gravenhorst (Marbled Salamander), which breeds in ephemeral wetlands much like Ambystoma maculatum Shaw (Spotted Salamander), was found at both sites, while the Spotted Salamander was not. The Marbled Salamander is reported as more tolerant of hotter, drier conditions than other ambystomatid species (Parmelee 1993). The woodland habitat at the Elon Site is a smaller fragment than at the Haw Site, and likely experiences higher temperatures and lower soil moisture levels. These conditions may explain the presence of the more tolerant Marbled Salamander at this smaller site. Another upland species, the White-spotted Slimy Salamander, was found only at the Haw Site. As a more terrestrial hardwood forest species, the White-spotted Slimy Salamander may be less sensitive to the availability of aquatic habitats as it is to historical and current woodland disturbances. However, Beamer and Lannoo (2005) reviewed literature on this species and report that it is relatively resilient to anthropogenic disturbances, and is often found in small woodland fragments. Therefore, we are uncertain why it was not found at the smaller Elon Site. Probable species and anecdotal observations We believe that the species inventory lists we accumulated from our study sites are typical of a suburban/agricultural/natural mosaic landscape within the Piedmont of North Carolina. We documented 28 of the 53 (52%) herpetofaunal species and 20 of the 32 (62%) non-volant mammal species we could potentially have encountered based on the distribution maps in Webster et al. (1985) and Beane et al. (2010). It is not surprising that certain species were found at the Haw Site and not the Elon Site, given the greater availability of 2012 J.M. Kapfer and D.J. Muñoz 79 water at the former (i.e., more and larger streams, as well as close vicinity to the Haw River). For example, Lutra canadensis Schreber (North American River Otter) is more often found in larger streams/rivers, such as those present at the Haw Site. We also observed a greater number of ephemeral wetlands, which retained water for longer periods of time, at the Haw Site. Although lowland habitat that superficially appeared suitable for ephemeral wetlands was present at the Elon Site, it did not hold water for as long of a period. This difference may explain why several amphibian species were found at the Haw Site, but not the Elon Site. We found the majority of species that we expected at the more disturbed Elon Site. There are several probable or possible species at the Haw Site which we did not detect, but would expect based on their occurrence at the nearby Elon Site. These include Microtus pennsylvanicus Ord (Meadow Vole), Diadophis punctatus L. (Ring-necked Snake), and Gastrophryne carolinensis Holbrook (Eastern Narrow-mouthed Toad). Mephitis mephitis Schreber (Striped Skunk) was detected at the Elon Site, but not the Haw Site, which we found somewhat strange. However, in July 2011, a roadkilled individual was observed just outside of the Haw Site’s boundaries, indicating they may have been present but undetected there. Aquatic turtles were under-represented in our samples due to the limited aquatic turtle trapping that was accomplished. Although aquatic turtle trapping at the Haw Site was conducted from May to July 2011 along the adjacent Haw River, only one Chelydra serpentina L. (Common Snapping Turtle) was captured. These surveys were terminated due to equipment theft. Based on range maps provided by Beane et al. (2010) and habitat on-site, we would expect Pseudemys concinna LeConte (River Cooter), Trachemys scripta Schoepff (Yellow-bellied Slider), Kinosternon subrubrum Lacèpède (Eastern Mud Turtle), Sternotherus odoratus Latreille in Sonnini and Latreille (Eastern Musk Turtle), and Chrysemys picta Schneider (Painted Turtle) to be present at the Haw Site. No suitable location for turtle traps existed in the smaller streams found at the Elon Site. We were surprised that several amphibian and reptile species found in the Piedmont went undetected at both sites. These include Lampropeltis getula L. (Eastern Kingsnake), Storeria occipitomaculata Storer (Red-bellied Snake), Virginia valeriae Baird and Girard (Smooth Earth Snake), and Notophthalmus viridescens Rafinesque (Eastern Newt). All of these species were observed in the Piedmont of North Carolina by Rice et al. (2001). Other relatively common snakes that we did not observe include Thamnophis sauritus L. (Eastern Ribbon Snake) and Regina septemvittata Say (Queen Snake). In general, these species are cryptic and may exist on-site, but in densities too low to be detected. A number of mammal species were surprisingly not detected at either site during our surveys: Tamias striatus L. (Eastern Chipmunk) and Sigmodon hipsidus Say and Ord (Hipsid Cotton Rat), both of which were found in the Piedmont by Kalcounis-Rueppel et al. (2007a). We also expected, for example, Sorex longirostris Bachman (Southeastern Shrew), and Mus musculus L. (House Mouse) based on the range maps in Webster et al. (1985). On 24 June 2011, an Ursus 80 Southeastern Naturalist Vol. 11, No. 1 americanus Pallas (American Black Bear) was witnessed concurrently by 10 county park and recreation department employees at the Haw Site. Attempts to obtain evidence of this species were mounted within 12–48 h of the observation. This effort included visual surveys for tracks and deployment of an additional camera trap at a unique location (baited with food lures). We continued this effort for four weeks without success. Based on the landscape associated with this site, the presence of U. americanus is reasonable for this location, although it is uncommon in the Piedmont of North Carolina. We also expected Lynx rufus Schreber (Bobcat) to be present at the Haw Site, and unsuccessful efforts were made to lure them to an additional unique camera trap with scent and food lures. We believe that, if present, they exist in densities too low to have been captured by camera traps. Although cat-like tracks were seen at this site on several occasions, they could not be definitively identified and may have been feral Domestic Cat. Further research and surveys that document the wild species in areas of the southern Piedmont are recommended. Acknowledgments R. Kirk (Elon University) conducted the GIS habitat assessments of both sites and reviewed historic aerial photos for land-use patterns. D. Vandermast (Elon University) assisted with outlining vegetation communities present at each site, and also provided information on historic land-use. B. Touchette (Elon University) provided assistance in designing and constructing survey equipment. Box-trap design was provided by R. Zappalorti (Herpetological Associates, Inc.). We thank B. Hagood, B. Baker, and R. Graves (Alamance County Recreation and Parks) for granting site access and logistical support. We thank Elon University’s Faculty Research and Development Program and Summer Undergraduate Research Experience Program for providing funds to purchase survey equipment. Elon University’s Center for the Advancement of Teaching and Learning also provided funding to purchase survey equipment. M. Kingston and the Department of Environmental Studies (Elon University) assisted with sequestering equipment funds. J. Balavender, K. Browning, J. Folkerts, O. Frey, S. Gerald, A. Keech, A. Maddalone, M. McGrath, K. Meredith, E. Neidhardt, R. Purnsley, M. Schriber, M. Strayer, and E. Winchester assisted with field collection of data. K. Rehrauer (Elon University) assisted with equipment purchase. All animals were treated humanely under the American Society of Ichthyologists and Herpetologists’ Guidelines for Use of Live Amphibians and Reptiles in Field and Laboratory Research and the Guidelines of the American Society of Mammalogists for the Use of Wild Mammals in Research. 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Inventory list, including abundances (n) and capture probabilities (CP; 95% confidence intervals) of each amphibian, reptile, and non-volant mammal species encountered or observed at two sites in the Piedmont of North Carolina (Alamance County). Common and scientific names taken from Webster et al. (1985), Feldhamer et al. (2003) and Beane et al. (2010). Habitat codes as follows: UF = Upland Forest, LF = Lowland Forest, GR = Grassland, RI = Riparian, ED = Edge. Elon site Haw site n CP Habitat n CP Habitat Class Amphibia Order Caudata Desmognathus fuscus Rafinesque 0 2 0.0002 (-0.0009, 0.0012) RI (Northern Dusky Salamander) Eurycea cirrigera Green 4 0.0007 (-0.0013, 0.0028) UF 9 0.0010 (-0.0013, 0.0033 RI (Southern Two-lined Salamander) Plethodon cylindraceus Harlan 0 50 0.5118 (0.3570, 0.665) UF (White-spotted Slimy Salamander) Ambystoma maculatum Shaw 0 4 0.0005 (-0.0012, 0.0021) LF (Spotted Salamander) Ambystoma opacum Gravenhorst 6 0.0005 (-0.0013, 0.0023) UF 13 0.1825 (0.0157, 0.3493) UF/LF (Marbled Salamander) Order Anura Anaxyrus = Bufo americanus Holbrook 26 0.2841 (0.1216, 0.4465) UF/LF 8 0.0010 (-0.0013, 0.0033) LF (American Toad) Anaxyrus = Bufo fowleri Hinckley 0 17 0.2372 (0.0834, 0.3910) LF (Fowler’s Toad) Gastrophryne carolinensis Holbrook 3 0.0004 (-0.0011, 0.0019) LF 0 (Eastern Narrow-mouthed Toad) Acris crepitans Baird 0 4 0.0005 (-0.0012, 0.0021) LF (Northern Cricket Frog) Lithobates = Rana palustris LeConte 1 0.0002 (-0.0010, 0.0013) RI 6 0.0010 (-0.0013, 0.0033) RI (Pickerel Frog) Lithobates = Rana clamitans Latreille in 2 0.0004 (-0.0011, 0.0019) RI/LF 11 0.0008 (-0.0013, 0.0030) LF Sonnini de Manoncourt and Latrielle (Green Frog) Lithobates = Rana catesbeiana Shaw 0 3 0.0005 (-0.0012, 0.0021) LF (Bullfrog) 86 Southeastern Naturalist Vol. 11, No. 1 Elon site Haw site n CP Habitat n CP Habitat Pseudacris crucifer Wied-Neuwied 3 0.0005 (-0.0013, 0.0023) LF 3 0.0005 (-0.0012, 0.0021) LF (Spring Peeper) Pseudacris feriarum Baird 0 3 0.0005 (-0.0012, 0.0021) LF (Upland Chorus frog) Hyla chrysoscelis Cope 3 0.0005 (-0.0013, 0.0023) LF 0 (Cope’s Gray Treefrog) Class Reptilia Suborder Serpentes Carphophis amoenus Say 28 0.4196 (0.2571, 0.5849) LF 15 0.1515 (-0.0537, 0.3568) UF (Eastern Worm Snake) Storeria dekayi Dekay 2 0.0004 (-0.0011, 0.0019) LF 0 (Brown Snake) Diadophis punctatus L. 1 0.0002 (-0.0010, 0.0013) UF 0 (Ring-necked Snake) Opheodrys aestivus L. 0 1 0.0002 (-0.0009, 0.0012) GR (Rough Green Snake) Thamnophis sirtalis L. 10 0.1970 (0.0183, 0.3756) GR 3 0.0005 (-0.0012, 0.0021) GR (Eastern Garter Snake) Nerodia sipedon L. 0 3 0.0003 (-0.0011, 0.0017) RI (Northern Water Snake) Coluber constrictor L. 8 0.2560 (0.0919, 0.4200) GR 9 0.0010 (-0.0013, 0.0033) GR (Black Racer) Elaphe = Pantherophis obsoletus Holbrook 4 0.0007 (-0.0013, 0.0028) GR 5 0.0008 (-0.0013, 0.0030) GR (Rat Snake) Agkistrodon contortix L. 2 0.0004 (-0.0011, 0.0019) UF/GR 7 0.0008 (-0.0013, 0.0030) GR (Copperhead) Suborder Lacertilia Scincella lateralis Say in James 2 0.0004 (-0.0011, 0.0019) GR 1 0.0002 (-0.0009, 0.0012) GR (Ground Skink) Plestiodon = Eumeces fasciatus L. 1 0.0002 (-0.0010, 0.0013) UF 3 0.0005 (-0.0012, 0.0021) UF (Five-lined Skink) Sceloporus undulatus Bosc and Daudin in 2 0.0002 (-0.0010, 0.0013) UF 0 Sonnini and Latreille (Eastern Fence Lizard) 2012 J.M. Kapfer and D.J. Muñoz 87 Elon site Haw site n CP Habitat n CP Habitat Order Testudines Chelydra serpentina L. 0 1 0.0002 (-0.0009, 0.0012) ED (Common Snapping Turtle) Terrapene carolina L. 22 0.3660 (0.2039, 0.5281) UF/LF 21 0.0028 (-0.0013, 0.0070) UF/LF (Eastern Box Turtle) Class Mammalia Order Marsupialia Didelphis virginiana Kerr 27 0.0010 (-0.0049, 0.0069) UF/LF 35 0.0417 (0.0261, 0.0572) UF/LF (Virginia Opossum) Order Insectivora Blarina carolinensis Bachman 5 0.0009 (-0.0014-0.0032) LF/GR 9 0.0008 (-0.0013-0.0030) GR (Southern Short-tailed Shrew) Order Rodentia Microtus pennsylvanicus Ord 1 0.0002 (-0.0010-0.0013) GR 0 (Meadow Vole) Microtus pinetorum LeConte 0 1 0.0002 (-0.0009-0.0012) LF (Woodland Vole) Reithrodontomys humulis Audubon & Bachman 3 0.0005 (-0.0013-0.0023) GR 4 0.0005 (-0.0012-0.0021) GR (Eastern Harvest Mouse) Peromyscus leucopus Rafinesque 8 0.0011 (-0.0014-0.0036) UF 42 0.5607 (0.4076-0.7138) UF (White-footed Mouse) Ochrotomys nuttalli Harlan 0 2 0.0003 (-0.0011-0.0017) UF (Golden Mouse) Zapus hudsonius Zimmermann 0 3 0.0003 (-0.0011-0.0017) GR (Meadow Jumping Mouse) Sciurus carolinensis Gmelin 39 0.0014 (-0.0048, 0.0076) LF/UF 34 0.0363 (0.0210, 0.0515) LF/UF (Eastern Gray Squirrel) Marmota monax L* 0 1 GR (Whistle Pig or Woodchuck) Castor canadensis Kuhl* 2 RI 3 RI (North American Beaver) 88 Southeastern Naturalist Vol. 11, No. 1 Elon site Haw site n CP Habitat n CP Habitat Order Lagomorpha Sylvilagus floridanus J.A. Allen 1 0.0001 (-0.0001, 0.0001) LF 3 less than 0.0001 (-0.0001, 0.0002) LF (Eastern Cottontail) Order Carnivora Procyon lotor L. 63 0.0084 (-0.0074, 0.0242) LF/RI 37 0.0094 (0.0017, 0.0174) LF/RI (Raccoon) Mephitis mephitis Schreber 1 0.0001 (-0.0001, 0.0001) GR 0 (Striped Skunk) Mustela vison Schreber* 0 8 RI (American Mink) Lutra canadensis Schreber* 0 11 RI (River Otter) Urocyon cinereoargenteus Schreber 34 0.0182 (0.0072, 0.0292) GR 4 0.0063 (-0.0020, 0.0146) LF (Gray Fox) Vulpes vulpes L. 1 0.0001 (-0.0001, 0.0001) LF 0 (Red Fox) Canis latrans Say 9 0.0002 (-0.0011, 0.0016) GR 9 <0.0001 (-0.0001, 0.0001) GR/UF (Coyote) Order Artiodactyla Odocoileus virginianus Zimmermann 433 0.116 (0.1011, 0.1312) UF/LF/GR/RI 933 0.1657 (0.1501, 0.1826) UF/LF/GR/RI (White-tailed Deer) Feral Species+ Canis familiaris L. 2 10 (Domestic Dog) Felis catus L. 1 0 (Domestic Cat) *Indicates a species that was only detected via tracks and/or scat or single visual observation for which capture probabilities were not calculated. +Indicates an animal not included in diversity measures.