Site by Bennett Web & Design Co.
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.
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 - email@example.com.
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.
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
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.
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.
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.
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
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).
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
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
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.
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.
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. Appropriate permits to live trap
wild amphibians, reptiles, and mammals were acquired from the North Carolina Wildlife
Atkeson, T.D., and A.S. Johnson. 1979. Succession of small mammals on pine plantations
in the Georgia Piedmont. American Midland Naturalist 101:385–392.
Barnosky, A.D., N. Matzke, S. Tomiya, G.O.U. Wogan, B. Swartz, T.B. Quental, C. Marshall,
J.L. McGuired, E.L. Lindsey, K.C. Maguire, B. Mersey, and E.A. Ferrer. 2011.
Has the earth’s sixth mass extinction already begun? Nature 471:51–57.
2012 J.M. Kapfer and D.J. Muñoz 81
Beamer, D.A., and M.J. Lannoo. 2005. Plethodon cylindraceus (Harlan, 1825[b]), Whitespotted
Slimy Salamander. Pp. 800–802, In M.J. Lannoo (Ed.). Amphibian Declines:
The Conservation and Status of United States Species. University of California Press,
Berkeley, CA. 1115 pp.
Beane, J.C., A.L. Braswell, J.C. Mitchell, W.M. Palmer, and J.R. Harrison III. 2010.
Amphibians and Reptiles of the Carolinas and Virginia. University of North Carolina
Press, Chapel Hill, NC. 288 pp.
Bekoff, M., and E.M. Gese. 2003. Coyote, Canis latrans. Pp. 467–481, In G.A. Feldhamer,
B.C. Thompson, and J.A. Chapman (Ed.). Wild Mammals of North America:
Biology, Management, and Conservation (Second Edition). Johns Hopkins University
Press, MD. 1216 pp.
Christopher, C.C., and G.W. Barrett. 2006. Coexistence of White-footed Mice (Peromyscus
leucopus) and Golden Mice (Ochrotomys nuttalli) in a southeastern forest.
Journal of Mammalogy 87:102–107.
Conroy, M.J., C.R. Allen, J.T. Peterson, L. Pritchard, Jr., and C.T. Moore. 2003. Landscape
change in the southern Piedmont: Challenges, solutions, and uncertainty across
scales. Conservation Ecology 8:3.
Cutler, T.L., and D.E. Swann. 1999. Using remote photography in wildlife ecology: A
review. Wildlife Society Bulletin 27:571–581.
Dorcas, M.E., S.J. Price, and G.E. Vaughan. 2006. Amphibians and reptiles of the Great
Falls Bypassed Reaches in South Carolina. Journal of the North Carolina Academy
of Science 122:1–9.
Dorcas, M.E., A.M. Domske, and G.E. Vaughan. 2010. A herpetofaunal inventory of
Lake Keowee and Lake Jocassee, South Carolina. Journal of the North Carolina
Academy of Science 126:88–97.
Feldhamer, G.A., B.C. Thompson, and J.A. Chapman. 2003. Wild Mammals of North
America: Biology, Management, and Conservation (Second Edition). Johns Hopkins
University Press, Baltimore, MD. 1216 pp.
Ford, W.M., M.A. Menzel, T.S. McCay, J.W. Gassett, and J. Laerm. 2000. Woodland
salamander and small-mammal responses to alternative silvicultural practices in the
southern Appalachians of North Carolina. Proceedings of the Annual Conference of
the Southeastern Association of Fish and Wildlife Agencies 54:241–250.
Foster, J., and M.S. Gaines. 1991. The effects of a successional habitat mosaic on a smallmammal
community. Ecology 72:1358–1373.
Fritzell, E.K., and K.J. Haroldson. 1982. Mammalian Species: Urocyon cinereoargenteus.
American Society of Mammalogists No. 189:1–8.
Hadidian, J., S. Prange, R. Rosatte, S.P.D. Riley, and S.D. Gehrt. 2010. Raccoons (Procyon
lotor). Pp. 35–47, In S.D. Gehrt, S.P.D. Riley, and B.L. Cypher (Eds.). Urban
Carnivores: Ecology, Conflict, and Conservation. Johns Hopkins University Press,
Baltimore, MD. 304 pp.
Heyer, W.R., M.A. Donelly, R.W. McDiarmid, L.C. Hayek, and M.S. Foster. 1994.
Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians.
Smithsonian Books, Washington, DC. 384 pp.
Hicks, N.G., and S.M. Pearson. 2003. Salamander diversity and abundance in forests
with alternative land-use histories in the southern Blue Ridge Mountains. Forest Ecology
and Management 177:117–130.
82 Southeastern Naturalist Vol. 11, No. 1
Hines, J.E. 2006. PRESENCE2-Software to estimate patch occupancy and related parameters.
USGS-PWRC, Reston, VA. Available online at http://www.mbr-pwrc.usgs.gov/
software/presence.html. Accessed on 29 May 2011.
Hutchens, S., and C. DePerno. 2009. Measuring species diversity to determine land-use
effects on reptile and amphibian assemblages. Amphibia-Reptilia 30:81–88.
Jost, L. 2010. The relation between evenness and diversity. Diversity 2:207–232.
Kalcounis-Rueppell, M.C., L. Shiflet, and M. Vidigni. 2007a. Non-volant mammal inventory
for Guilford Courthouse National Military Park (GUCO) within the Cumberland
Piedmont Network. Prepared for the National Park Service, Guilford Courthouse
National Military Park (catalogue # GUCO12187).
Kalcounis-Rueppell, M.C., V.H. Payne, S.R. Huff, and A.L. Boyko. 2007b. Effects of
wastewater treatment plant effluent on bat foraging ecology in an urban stream system.
Biological Conservation 138:120–130.
Kaminski, J.A., M.L. Davis, M. Kelly, and P.D. Keyser. 2007. Disturbance effects on
small mammal species in a managed Appalachian forest. American Midland Naturalist
Karns, D.R. 1986. Field herpetology: Methods for the study of amphibians and reptiles
in Minnesota. Occasional Paper No. 18. James Ford Bell Museum of Natural History,
University of Minnesota, Minneapolis, MN. 88 pp.
Kelly, M.J. 2008. Design, evaluate, refine: Camera-trap studies for elusive species. Animal
Krebs, C.J. 1998. Ecological Methodology (2nd Edition). Benjamin Cummings, Menlo
Park, CA. 624 pp.
Kremen, C. 1994. Biological inventory using target taxa: A case study of the butterflies
of Madagascar. Ecological Applications 4:407–422.
Lackey, J.A., D.G. Huckaby, and B.G. Ormiston. 1985. Mammalian Species: Peromyscus
leucopus. American Society of Mammalogists 247:1–10.
Matthews, W.J. 1990. Spatial and temporal variation in fishes of riffle habitats: A comparison
of analytical approaches for the Roanoke River. American Midland Naturalist
Metts, B.S., J.D. Lanham, and K.R. Russell. 2001. Evaluation of herpetofaunal communities
on upland streams and Beaver-impounded streams in the upper Piedmont of
South Carolina. American Midland Naturalist 145:54–65.
Meyers, J.M., and D.A. Pike. 2006. Herpetofaunal diversity of Alligator River National
Wildlife Refuge, North Carolina. Southeastern Naturalist 5:235–252.
McKinney, M.L. 2006. Urbanization as a major cause of biotic homogenization. Biological
Mitchell, J., and W. Gibbons. 2010. Salamanders of the Southeast. University of Georgia
Press, Athens, GA. 336 pp.
Mitchell, M.S., K.S. Karriker, E.J. Jones, and R.A. Lancia. 1995. Small-mammal communities
associated with pine plantation management of pocosins. Journal of Wildlife
O’Connell, A.F., J.D. Nichols, and K.U. Karranth. 2011. Camera Traps in Animal Ecology:
Methods and Analyses. Springer Publishing, New York, NY. 280 pp.
Osbourne, J.D., J.T. Anderson, and A.B. Spurgeon. 2005. Effects of habitat on small-mammal
diversity and abundance in West Virginia. Wildlife Society Bulletin 33:814–822.
2012 J.M. Kapfer and D.J. Muñoz 83
Pagels, J.F., S.Y. Erdle, K.L. Uthus, and J.C. Mitchell. 1992. Small-mammal diversity in
forested and clearcut habitats in the Virginia Piedmont. Virginia Journal of Science
Parmelee, J.R. 1993. Microhabitat segregation and spatial relationships among four species
of mole salamander (genus Ambystoma). Occasional Papers of the Museum of
Natural History, Number 160, University of Kansas, Lawrence, KS.
Pearson, P.G., 1953. A field study of Peromyscus populations in Gulf Hammock, Florida.
Price, S.J., M.E. Dorcas, A.L. Gallant, R.W. Klaver, and J.D. Willson. 2006. Three
decades of urbanization: Estimating the impact of land-cover change on stream salamander
populations. Biological Conservation 133:436–441.
Price, S.J., K.K. Cecala, R.A. Browne, and M.E. Dorcas. 2010. Effects of urbanization
on occupancy of stream salamanders. Conservation Biology 25:547–555.
Primack, R.B. 2010. Essentials of Conservation Biology, Fifth Edition. Sinauer Associates,
Inc. Sunderland, MA. 601 pp.
Rice, A.N., T.L. Roberts IV, J.G. Pritchard, and M.E. Dorcas. 2001. Historical trends
and perceptions of amphibian and reptile diversity in the western Piedmont of North
Carolina. The Journal of the Elisha Mitchell Scientific Society 117:264–273.
Roseberry, J.L., and A. Woolf. 1998. Habitat-population density relationships for Whitetailed
Deer in Illinois. Wildlife Society Bulletin 26:252–258.
Royle, J.A., and J.D. Nichols. 2003. Estimating abundance from repeated presenceabsence
data or point counts. Ecology 84:777–790.
Ryan, T.J., T. Philippi, Y.A. Leiden, M.E. Dorcas, T.B. Wigley, and J.W. Gibbons. 2002.
Monitoring herpetofauna in a managed forest landscape: Effects of habitat types and
census techniques. Forest Ecology and Management 167:83–90.
Schemnitz, S.D. 2005. Capturing and handling wild animals. Pp. 239–285, In C. Braun
(Ed.). Techniques for Wildlife Investigations and Management, Sixth Edition. The
Wildlife Society, Bethesda, MD. 974 pp.
Spira, T.P. 2011. Wildflowers and Plant Communities of the Southern Appalachian Mountains
and Piedmont. University of North Carolina Press, Chapel Hill, NC. 584 pp.
Todd, B.D., C.T. Winne, J.D. Willson, and J. Whitfield Gibbons. 2007. Getting the drift:
Examining the effects of timing, trap type, and taxon on herpetofaunal drift-fence
surveys. American Midland Naturalist 158:292–305.
Todd, M.J., R.R. Cocklin, and M.E. Dorcas. 2003. Temporal and spatial variation in
anuran calling activity in the western Piedmont of North Carolina. Journal of the
North Carolina Academy of Science. 119:103–110.
Tuberville, T.D., J.D. Willson, M.E. Dorcas, and J.W. Gibbons. 2005. Herpetofaunal species
richness of southeastern national parks. Southeastern Naturalist 4:537–569.
United States Census Bureau. 2010a. Estimates of resident population change for the
United States: Regions, states, and Puerto Rico and region and state rankings: July 1,
2008 to July 1, 2009. Available online at http://www.census.gov/popest/states/NSTpop-
chg.html. Accessed on 27 June 2011.
United States Census Bureau. 2010b. State and county quick facts: North Carolina.
Available online at http://quickfacts.census.gov/qfd/states/37000.html. Accessed on
27 June 2011.
Wear, D.N., and J.G. Greis. 2001. The southern forest research assessment: Draft summary
report. US Forest Service, Asheville, NC.
84 Southeastern Naturalist Vol. 11, No. 1
Webster, W.D. 2005. The mammals of the Great Falls Bypassed Reaches (Great Falls-
Dearborn Development) in South Carolina. Final Report to Duke Power Company.
Available online at http://www.duke-energy.com/pdfs/Great__Falls_Bypass_mammals.
pdf. Accessed 17 July 2011.
Webster, W.D., J.F. Parnell, and W.G. Biggs, Jr. 1985. Mammals of the Carolinas, Virginia,
and Maryland. University of North Carolina Press, Chapel Hill, NC. 272 pp.
Welsh, H.H., Jr., and S. Droege. 2001. A case for using plethodonid salamanders for
monitoring biodiversity and ecosystem integrity of North American forests. Conservation
Whitaker, J.O. 1972. Mammalian Species: Zapus hudsonius. American Society of Mammalogists
Wilder, S.M., A.M. Abtahi, and D.B. Meikle. 2005. The effects of forest fragmentation
on densities of White-footed Mice (Peromyscus leucopus) during the winter. American
Midland Naturalist 153:71–79.
Willson, J.D., and M.E. Dorcas. 2004. Aspects of the ecology of small fossorial snakes in
the western Piedmont of North Carolina. Southeastern Naturalist 3:1–12.
Wilson, D.E., F.R. Cole, J.D. Nichils, R. Rudran, and M.S. Foster. 1996. Measuring
and Monitoring Biological Diversity: Standard Methods for Mammals. Smithsonian
Books, Washington, DC. 440 pp.
2012 J.M. Kapfer and D.J. Muñoz 85
Appendix 1. 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
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
Ambystoma opacum Gravenhorst 6 0.0005 (-0.0013, 0.0023) UF 13 0.1825 (0.0157, 0.3493) UF/LF
Anaxyrus = Bufo americanus Holbrook 26 0.2841 (0.1216, 0.4465) UF/LF 8 0.0010 (-0.0013, 0.0033) LF
Anaxyrus = Bufo fowleri Hinckley 0 17 0.2372 (0.0834, 0.3910) LF
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
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
Lithobates = Rana catesbeiana Shaw 0 3 0.0005 (-0.0012, 0.0021) LF
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
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)
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
Diadophis punctatus L. 1 0.0002 (-0.0010, 0.0013) UF 0
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
Elaphe = Pantherophis obsoletus Holbrook 4 0.0007 (-0.0013, 0.0028) GR 5 0.0008 (-0.0013, 0.0030) GR
Agkistrodon contortix L. 2 0.0004 (-0.0011, 0.0019) UF/GR 7 0.0008 (-0.0013, 0.0030) GR
Scincella lateralis Say in James 2 0.0004 (-0.0011, 0.0019) GR 1 0.0002 (-0.0009, 0.0012) GR
Plestiodon = Eumeces fasciatus L. 1 0.0002 (-0.0010, 0.0013) UF 3 0.0005 (-0.0012, 0.0021) UF
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
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)
Didelphis virginiana Kerr 27 0.0010 (-0.0049, 0.0069) UF/LF 35 0.0417 (0.0261, 0.0572) UF/LF
Blarina carolinensis Bachman 5 0.0009 (-0.0014-0.0032) LF/GR 9 0.0008 (-0.0013-0.0030) GR
(Southern Short-tailed Shrew)
Microtus pennsylvanicus Ord 1 0.0002 (-0.0010-0.0013) GR 0
Microtus pinetorum LeConte 0 1 0.0002 (-0.0009-0.0012) LF
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
Ochrotomys nuttalli Harlan 0 2 0.0003 (-0.0011-0.0017) UF
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
Sylvilagus floridanus J.A. Allen 1 0.0001 (-0.0001, 0.0001) LF 3 less than 0.0001 (-0.0001, 0.0002) LF
Procyon lotor L. 63 0.0084 (-0.0074, 0.0242) LF/RI 37 0.0094 (0.0017, 0.0174) LF/RI
Mephitis mephitis Schreber 1 0.0001 (-0.0001, 0.0001) GR 0
Mustela vison Schreber* 0 8 RI
Lutra canadensis Schreber* 0 11 RI
Urocyon cinereoargenteus Schreber 34 0.0182 (0.0072, 0.0292) GR 4 0.0063 (-0.0020, 0.0146) LF
Vulpes vulpes L. 1 0.0001 (-0.0001, 0.0001) LF 0
Canis latrans Say 9 0.0002 (-0.0011, 0.0016) GR 9 <0.0001 (-0.0001, 0.0001) GR/UF
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
Canis familiaris L. 2 10
Felis catus L. 1 0
*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.