An Overlooked Hotspot? Rapid Biodiversity Assessment
Reveals a Region of Exceptional Herpetofaunal Richness
in the Southeastern United States
Sean P. Graham, David A. Steen, Kerry T. Nelson, Andrew M. Durso,
and John C. Maerz
Southeastern Naturalist, Volume 9, Issue 1 (2010): 19–34
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2010 SOUTHEASTERN NATURALIST 9(1):19–34
An Overlooked Hotspot? Rapid Biodiversity Assessment
Reveals a Region of Exceptional Herpetofaunal Richness
in the Southeastern United States
Sean P. Graham1,*, David A. Steen1, Kerry T. Nelson2, Andrew M. Durso3,
and John C. Maerz2
Abstract - We conducted a competitive bioblitz survey in four Georgia counties
to raise awareness of a unique and species rich herpetofauna in the Pine Mountain/
Fall Line Sandhills Region of Georgia, and compared documented species of these
counties to other herpetofaunas of the southeast that have known high richness and/
or were subject to thorough collection efforts. Our results demonstrate the efficacy
of bioblitzes for documenting large numbers of species in a limited amount of time
(62 amphibian and reptile species in only seven days, including 36 new county records
and documentation of three protected species). Compared to areas of similar
size, this area is among the most species-rich herpetofaunas in North America north
of Mexico, with only three areas having higher documented richness. However, all
areas we compared our site to have experienced much higher collection effort and
contain much larger tracts of protected land. Thus, our data suggest the Pine Mountain/
Fall Line Sandhills region is among the most important regions for amphibian
and reptile conservation in North America.
Biodiversity hotspots—areas undergoing extensive habitat loss
characterized by high species richness, diversity, and/or endemism—
have garnered great attention from ecologists, land managers, and conservationists
as interesting areas for research and practical targets for
conservation (Myers et al. 2000). A range of factors influence patterns of
species richness within a given area, including local environmental variables
(Qian et al. 2007), historical climate (Araú jo et al. 2008), latitude
(Pianka 1966), topography and/or habitat heterogeneity (Kerr and Packer
1997), and productivity (Rodrí guez et al. 2005). The first step in identifying
and understanding hotspots is to accurately measure species distributions
and richness. Not surprisingly, attention to hotspots is biased toward
easily sampled taxa (e.g., plants; Myers 1988, 1990). Unfortunately,
biodiversity hotspots do not always show great congruence among taxa,
so characterizations of one set of organisms may not apply to other taxa
(e.g., Daniels 1992, Grenyer et al. 2006). This lack of congruence presents
a unique challenge in identifying biodiversity hotspots for cryptic
taxa that are difficult to detect.
1Auburn University Department of Biological Sciences, 331 Funchess Hall, Auburn
University, AL 36849. 2Warnell School of Forestry and Natural Resources, University
of Georgia, Athens, GA 30602. 3Odum School of Ecology, University of Georgia,
Athens, GA 30602. *Corresponding author - email@example.com.
20 Southeastern Naturalist Vol. 9, No. 1
Amphibians and reptiles (hereafter, “herpetofauna”) are examples of taxa
that can be difficult to systematically inventory (Heyer et al. 1994) and therefore
may have regional hotspots that remain undocumented. Identification of
herpetofaunal hotspots may require opportunistic or targeted inventories and
long-term accumulation of records. For example, it took 45 years of incidental
and opportunistic encounters to identify areas within the Savannah River
Site (SRS) that contain as many as 90 reptile and amphibian species, and the
SRS is arguably one of the most intensively studied areas for herpetofauna
in the world (Gibbons et al. 1997). Even still, one amphibian species escaped
detection for over 50 years until only recently being discovered on the site
The southeastern United States is the center of herpetofaunal biodiversity
in North America (north of Mexico), containing approximately
half of its known species of amphibians and reptiles (Conant and Collins
1998, Gibbons and Buhlmann 2001, Tuberville et al. 2005). Therefore,
the location of the most species rich herpetofaunal assemblage in North
America presumably occurs in the southeastern United States, and it has
been suggested that the Florida Panhandle has the most amphibian and
reptile species for its size compared to any other region in North America
(Blaustein 2008). However, due to the inherent difficulties involved in
sampling these animals (Heyer et al. 1994), it is possible that other areas
with equal or higher species richness exist.
The Fall Line along the Gulf and Atlantic Coasts of the United States has
particularly high species richness due to the intermingling of species typical
of the Coastal Plain and those restricted to other physiographic provinces
(e.g., Piedmont, Ridge and Valley, etc.; Mount 1975). In particular, one
section of the Georgia Fall Line appears to have notable species richness
(see Griffith et al. 2001, Wharton 1978). In this zone (the Pine Mountain
Ecoregion and nearby Fall Line Sandhills), certain northern/montane-associated
species (e.g., Rana sylvatica LeConte [Wood Frog], Gyrinophilus
porphyriticus Green [Spring Salamander]) reach the southernmost termini
of their ranges (Conant and Collins 1998, Jensen et al. 2008), while Coastal
Plain-associated species (e.g., Rana capito LeConte [Gopher Frog], Crotalus
adamanteus Palisot de Beauvois [Eastern Diamondback Rattlesnake],
and Notophthalmus perstriatus Bishop [Striped Newt]) occur in some of
their inland-most populations (Jensen et al. 2008). Surprisingly, this area
has not been subject to intensive study, although its significance relative
to vertebrates has been briefl y noted (e.g., Wharton 1978), and a botanical
study has been conducted, documenting similar patterns of biogeographic
intermingling of plant species (Jones 1974).
This paper presents the findings of a bioblitz to raise awareness of this
potentially important area of reptile and amphibian biodiversity. Bioblitz
competitions are events in which teams of biologists compete to document
and voucher the most species in a particular area (Graham et al. 2007a; Graham
et al., in press). Our goals were to: 1) document new county records to
2010 S.P. Graham, D.A. Steen, K.T. Nelson, A.M. Durso, and J.C. Maerz 21
supplement the list of known species and demonstrate the incompleteness
of sampling for this region, 2) document the presence or persistence of rare
and protected species, and 3) compare the species list for this region to wellstudied
herpetofaunas in the southeast known for their high diversity.
The core area of our survey included the four Georgia counties of Talbot,
Taylor, Marion, and Schley, although participants were encouraged to
locate new records in adjacent counties as well. Many terrestrial and aquatic
environments are found in this region; for complete descriptions of these
plant and animal communities, consult Wharton (1978). The Fall Line—the
physiographic boundary between the Piedmont and Coastal Plain—transects
southern Talbot and northern Taylor counties (Fig. 1). Areas north of this
boundary contain heavy, clay soils derived from crystalline metamorphic
rocks, whereas deep sands of the Coastal Plain are found south of it (Griffith
et. al. 2001, Wharton 1978). Southern portions of the Pine Mountain ridges
terminate in Talbot County (Fig. 1), and are composed of ancient (e.g., >1
billion years old) “basement” (e.g., continental shield) metamorphic rocks
(Steltenpohl et al. 2008). These ridges are topped with xeric mountain
Longleaf Pine-Blackjack Oak forests, and their north-facing slopes contain
mesic hardwood forests. Streams of this area often have high gradients and
contain larger, cobble substrates than those found elsewhere in the Piedmont
(Griffith et al. 2001). Typical oak-hickory assemblages are found in the Piedmont
hills surrounding these ridges.
The Fall Line Sandhills in this area once contained excellent tracts of
xeric Longleaf Pine-Turkey Oak forests which are now heavily fragmented,
with much of the original Pinus palustris Mill (Longleaf Pine) removed
(Wharton 1978). Within these sand ridges are black-water creeks and a few
remaining isolated wetlands. Whitewater Creek, a unique, clear, black-water
creek, is one of the sole locations in Georgia for Chamaecyparis thyoides L.
(Atlantic White Cedar) and many other plants (Patrick et al. 1995). The Flint
River is the largest stream fl owing through this area and contains numerous
shoals, riffl es, and gorges in its Piedmont section. Its fl oodplain widens extensively
after entering the Coastal Plain to form Magnolia Swamp (Fig. 1).
Most of the remaining tracts of natural forests and wetlands are in private
ownership (including Magnolia Swamp), and are surrounded by agriculture,
old fields, pine plantations, and rural development.
Twenty-five individuals in two teams participated in the survey. However,
daily effort ranged widely. The two teams competed to find undocumented
species throughout the region, seeking suitable habitat (with landowner’s
permission on private tracts), conducting visual encounter surveys, and thoroughly
turning cover objects searching for herpetofauna. Limited trapping
22 Southeastern Naturalist Vol. 9, No. 1
(using hoop and minnow traps) was conducted in ponds and creeks. Our survey
was divided into spring (21–23 March 2008) and fall (8–10 October 2008
and 26 Oct 2008) portions, a strategy previously demonstrated to maximize
results by accommodating diverse reptile and amphibian activity seasons
(Graham et al., in press; Todd et al. 2007). New records were vouchered as
Figure 1. Map of area surveyed for amphibians and reptiles over seven total days
in March and October 2008. Insert: four county center of our survey area, with key
physiographic features indicated.
2010 S.P. Graham, D.A. Steen, K.T. Nelson, A.M. Durso, and J.C. Maerz 23
digital photographs or specimens, verified by experts (see Appendix 1), and
deposited in the Auburn University Herpetological Collections (AUM).
Comparison to other southeastern US herpetofaunas with high species
We compiled lists of herpetofaunas in the Southeast from published
reports combined with museum records. We chose areas of known high species
richness based on overlapping ranges in published range maps (Mount
1975; Conant and Collins, 1998; Jensen et al., 2008), our own experience
with this region, and suggestions in Gibbons et al. (1997) and Blaustein
(2008). We focused on regions with 80 to >100 potential species based upon
overlapping ranges (Conant and Collins 1998), then chose localities within
these zones that have been well-studied and have published species lists (e.g,
Apalachicola National Forest [Means 1976], Savannah River Site [Gibbons
et al. 1997], Ichauway [Smith et al. 2006]). Our method resulted in 12 locations
of high species richness, varying area, and variable collection effort, to
which we compared our area of interest. It is likely that these represent some
of the most species rich herpetofaunas (of similar size) in North America
(Blaustein 2008). These areas ranged from small national forests (Tuskegee
National Forest, Macon County, AL; 45 km2 ) to large counties (Mobile and
Baldwin County, AL; 9508 km2). We chose these areas since they are high
in amphibian and reptile species richness and also fairly well studied (e.g.,
they have a long history of sampling due to repeated visits by university
biologists), and thus they have been sufficiently sampled. Since our goal
was not to provide an exhaustive analysis of species richness patterns in the
Southeast, we argue this is an appropriate approach and a heuristic exercise
to put our focal region in perspective.
For each region, lists included only native species that can be unequivocally
assigned to a species using morphology alone, and lists included
members of species complexes separable by molecular characters alone only
once (e.g., the Plethodon glutinosus Green [Slimy Salamander] complex).
This was a conservative approach that likely underestimated the species pool
for our own area by at least three species (i.e., the ranges of three members
of the P. glutinosus complex overlap in our study area [Jensen et al. 2008],
as well as the Elaphe allegheniensis Holbrook [Eastern Ratsnake]/Elaphe
spiloides Duméril, Bibron, and Duméril [Grey Ratsnake] contact zone
[Burbrink et al. 2000]). We did not include undescribed species known or
rumored to occur in an area. We included all species reported within the past
100 years as still present in each area.
Species richness is likely related to both study area size and collection
effort. To determine if species richness was associated with the area of
each location, we used linear regression, and calculated the per-area species
richness of each location. We then took residuals from this analysis
to rank locations above and below the mean richness of our sample of
sites. Since collection effort varied between localities, we determined
a qualitative estimate of effort for each area prior to this study, with no
24 Southeastern Naturalist Vol. 9, No. 1
surveys and/or collection efforts from university herpetologists considered
to be a “low” collection effort, one herpetofaunal survey and/or collection
effort from university herpetologists considered to be a “moderate” collection
effort, and localities with long-term collection data from university
herpetologists and/or numerous surveys considered to be “high” collection
efforts. Similarly, we qualitatively estimated the amount of protection for
each locality, with national forests, parks, and wildlife refuges considered
“highly” protected, military reservations and other federal properties (e.g.,
Department of Energy) considered “moderately” protected, and areas with
mostly private lands (e.g., counties) with “low” protection. We used SPSS
for statistical analyses, with α set at 0.05.
Results and Discussion
In seven days (disjunct between spring and fall periods), we documented
62 total species of amphibians and reptiles, including 36 new county
records (for 5 counties; Appendix 1), surpassing totals achieved during
previous bioblitz competitions conducted in other areas (Graham et al.
2007a; Graham et al., in press). Twenty-three of the new records we report
were from our core area (the rest from adjacent counties; see Appendix 1),
with 54 species documented from Talbot, Taylor, Schley, and Marion counties.
By comparison, it took several decades to document 60 reptile and
amphibian species in most areas on the Savannah River Site (SRS), and we
documented approximately 2/3 the total number of species known from the
SRS (Gibbons et al. 1997) in seven days using limited trapping techniques.
However, it is important to mention that a more concerted effort using a
single drift fence documented 59 species at the SRS in just months (Todd
et al. 2007). A recent survey of species richness patterns in southeastern
national parks documented a maximum park species richness of 64, with
25 new county records after two years of detailed surveys using a variety
of standard methods (Tuberville et al. 2005). Our results illustrate the value
and utility of bioblitz competitions for generating baseline species lists in
general, as well as the extraordinary richness of the herpetofauna in this
area. Combined with previously documented species for these counties
(Jensen et al. 2008), our results indicate 104 documented amphibians and
reptiles for this region.
The comparison among localities in the Southeast demonstrated a positive
correlation between locality size and species richness (R2 = 0.349, F = 5.351,
P = 0.043), with our area exhibiting richness comparable to sites noted for
their herpetofaunal abundance and/or collection effort (Fig. 2, Table 1). Our
area had the fourth highest residual richness of our sample of localities (Table
1). However, this can be considered one of the first surveys of this area, and
thus collection effort has presumably been much lower for this area than any
of the other sites we compared it to (Table 1). To compensate for this lack of
collection effort, we analyzed predicted species richness for our area based
on range maps in Jensen et al. (2008). Including these species would raise our
2010 S.P. Graham, D.A. Steen, K.T. Nelson, A.M. Durso, and J.C. Maerz 25
pool to 111 species and result in the highest residual richness of our sample
(Fig. 2). It is likely that at least nine more species await documentation from
this region, such as the widespread Bufo americanus Holbrook (American
Toad), Plethodon serratus Grobman (Southern Redback Salamander), and Regina
septemvittata Say (Queen Snake), as well as the more secretive Farancia
erytrogramma Palisot de Beauvois (Rainbow Snake), Ophisaurus attenuatus
Cope (Slender Glass Lizard), and others. If these are documented in our area,
the Pine Mountain/Fall Line Sandhills region may possibly deserve status as
the most species-rich herpetofauna in North America.
A brief consideration of other herpetofaunas in the United States
demonstrates the high species richness of this region. For example, the
3242-km2 Big Bend National Park, which boasts more documented birds,
bats, and cacti (National Park Service 2009) than any other national park, has
67 documented amphibians and reptiles (56 of which are reptiles; National
Park Service 2009). The 370-km2 Saguaro National Park has only 54 species
(National Park Service 2009). Although the southwestern United States is
known for its reptile (especially lizard) diversity (Ricketts et al. 1999), the
Pine Mountain/Fall Line Sandhills region, with 60 species, still outranks
the reptile total for both of these southwestern national parks.
Figure 2. Relationship between amphibian and reptile species richness and size of
study area for regions with documented high species richness/and or extensive collection
effort in the southeastern United States.
26 Southeastern Naturalist Vol. 9, No. 1
Great Smoky Mountains National Park is heralded for its amphibian diversity
with 44 documented species (Dodd 2004, Tilley and Huheey 2001),
yet our focal area has the same number of documented amphibians. In sum,
one could combine the reptile species richness of Big Bend National Park
and the amphibian species richness of Great Smoky Mountain National Park,
and the herpetofauna species list would still be surpassed by our study area.
The herpetofaunal species richness observed at our study area and elsewhere
in the southeastern United States approaches the richness documented for
the tropics (Zug 1993:286–287).
Unfortunately, construction of a species-accumulation curve was not
practical due to daily fl uctuations in sampling effort (Gotelli and Colwell
2001), and therefore we cannot estimate how many total species may be
present. However, the number of new records we documented confirms that
sampling has been incomplete in this region, and that increased sampling
would likely result in documentation of additional species in the study area.
Recent discoveries of range extensions for Rana sylvatica (Graham et al.
2007b), and cryptic species such as Micrurus fulvius L. (Coral Snake) (Klaus
and Jensen 2009) in Talbot County are illustrative examples. While this paper
was under review, one of us (S.P. Graham) discovered a Desmognathus
aeneus Brown and Bishop (Seepage Salamander) on Rockhouse Mountain
in Talbot County (Graham 2009). This is yet another species with montane
affinities discovered in the Pine Mountain ecoregion. This species was not
included in our analysis.
Table 1. Species richness patterns in selected southeastern sites with high species richness. Sites
are represented in ascending order of residual richness. NF = National Forest, NWR = National
Wildlife Refuge, and AFB = Air Force Base. SR = species richness, RR = residual richness,
and CE = collection effort, AP = area preserved. H = high collection effort/preservation, M =
moderate collection effort/preservation, L = low collection effort/preservation.
Site (km2) SR Species/area RR CE AP Source
Apalachicola NF, FL 2286 106 0.04636920 1.21 M H Means 1976
St. Marks NWR, FL 280 99 0.35357143 0.81 H H USFWS 1998b
Savannah River Site, SC 803 99 0.12328767 0.66 H M Gibbons et al. 1997
Taylor, Talbot, Schley, 3379 104 0.03077834 0.63 L L This study,
and Marion counties, GA Jensen et al. 2008
Eglin AFB, FL 1875 100 0.05333333 0.49 M M Printiss and Hipes
Conecuh NF, AL 340 96 0.28235294 0.37 H H Guyer et al. 2007,
Fort Stewart, GA 1100 97 0.08818182 0.30 H M Stevenson 1999
Okefenokee NWR, GA 1627 98 0.06023356 0.28 H H USFWS 1998a
Mobile and Baldwin 9508 110 0.01156920 -0.67 M L Mount 1975;
Counties, AL Palmer 1987;
Carey 1984, 1985
Tuskegee NF, AL 45 86 1.91111111 -0.97 H H AUM records
Ichauway, GA 117 84 0.71794872 -1.28 H H Smith et al. 2006
Great Smoky Mountains 2108 81 0.03842505 -2.26 H H Tilley and Huheey
National Park, TN and NC 2001
2010 S.P. Graham, D.A. Steen, K.T. Nelson, A.M. Durso, and J.C. Maerz 27
In addition to increasing the documented species pool for counties in
this region, we documented the persistence of three rare/protected species.
Gopherus polyphemus Daudin (Gopher Tortoise) were observed at two sites,
a Macrochelys temminckii Gray (Alligator Snapping Turtle) was trapped in
a farm pond, and Gopher Frog egg masses were located at a historic locality
not resurveyed for this species since 1975 (based on Auburn University
Two historical and topographical factors appear to be responsible for
the high species richness in the region. First, the Apalachicola drainage is
a known biogeographic corridor and refugium that has facilitated historical
migrations during recent glacial advances and retreats (Blaustein 2008).
The Apalachicola Ravines region of the Florida Panhandle is a recognized
global hotspot with many endemics thought to have relict distributions (e.g.,
the critically endangered, Torreya taxifolia Arn [Florida Torreya]; Blaustein
2008). The Pine Mountain/Fall Line Sandhills area is also bisected by Apalachicola
drainages, and shares some of the same endemics with the ravines
further south (e.g., Silene polypetala Walter [Eastern Fringed Catchfl y]).
Therefore, this region probably harbored fl ora and fauna (including herpetofauna)
from higher latitudes during climatic fl uctuations, and they likely
migrated through the Apalachicola corridor. In addition, the Pine Mountain
Ridges trend east–west rather than north–south as most Appalachian ranges
do—similar topography has been suggested to have increased diversity and
endemism in Asian forests relative to North American ones during the Pleistocene
(Qian and Ricklefs 1999).
Second, the nearness of the Pine Mountain ecoregion to a rich and
almost complete Coastal Plain fauna results in the proximity of species
usually found much farther apart. Thus, this region represents an ecotone
at the ecoregion scale. Almost all southeastern habitat types (with the exception
of caves, high elevation cove/boreal forests, and coastal/marine
ecosystems) are available. The southernmost breeding population of Wood
Frogs (Graham et al. 2007b) occurs less than 30 km from a population of Gopher
Frogs and Striped Newts (Jensen 2000; Jensen and Klaus 2004), both
Coastal Plain endemics. We documented Spring Salamanders and Siren
lacertina L. (Greater Siren) during the same survey in nearly adjacent
counties (≈80 km apart). Future considerations of hotspots—regardless
of taxon of interest—should scrutinize border areas between distinctive
ecoregions. Additional studies are needed to determine if other taxa exhibit
similarly high richness in this region.
Areas such as Apalachicola National Forest, Great Smoky Mountains
National Park, and the Savannah River Site are known for their high diversity,
and their protection as public lands has maintained historical levels
of richness. Unfortunately, our focal region (especially in areas below the
Fall Line) has not enjoyed this advantage. Although this region has high local
herpetofaunal species richness, little of this area is protected as public
land, and habitat in the Fall Line Sandhills is generally degraded for much
28 Southeastern Naturalist Vol. 9, No. 1
wildlife. Most of the area is heavily fragmented and under intensive agriculture
or silviculture (see Wharton 1978:183). The region may still be suitable
for the acquisition and restoration of large areas as conservation easements.
Fortunately, a nearby military base (Fort Benning) utilizes conservation
management, parts of the Pine Mountain Ridges are protected as state lands
(FDR State Park in Harris and Meriwether Counties, Big Lazar Creek WMA
in Talbot County, and Sprewell Bluff State Park in Upson County), and the
recent designation of Fall Line Sandhills Natural Area by the Georgia Department
of Natural Resources (which protects an isolated wetland used by
rare Coastal Plain amphibians) is progress toward this goal.
The number of endemic species in the area and the extent of habitat loss
may be insignificant on a large scale, not qualifying this region as a global
hotspot (Myers et al. 2000). However, on a more local scale, we suggest that
its high species richness and diversity, coupled with heavy habitat modification
and lack of protected land in the area, may qualify the Pine Mountain/
Fall Line Sandhills ecotone as one of the most important areas for herpetofaunal
conservation within the United States and Canada.
Numerous individuals collected data associated with this project. We thank D.B.
Means, D. Printiss, and D. Stevenson for their help supplying references and data for
the herpetofaunas listed in Table 1. We thank C. Guyer, J. Jensen, C. Camp, and J.
Godwin for their time verifying records. We thank the editorial board of the Southeastern
Naturalist, Brian Todd, and an anonymous reviewer for their suggestions
which considerably improved this document. S.P. Graham is supported by NIH grant
# R01-A149724 to T. Unnasch under GA collection permit # 29-WTN-07-41 and 29-
Araú jo, M., D. Nogú es-Bravo, J.A.F. Diniz-Filho, A.M. Haywood, P.J. Valdes, and
C. Rahbek. 2008. Quaternary climate changes explain diversity among reptiles
and amphibians. Ecography 31:8–15.
Blaustein, R.J. 2008. Biodiversity hotspot: The Florida Panhandle. BioScience.
Burbrink, F.T., R. Lawson, and J.B. Slowinski. 2000. Mitochondrial DNA phylogeography
of the polytypic North American Rat Snake (Elaphe obsoleta): A
critique of the subspecies concept. Evolution 54:2107–2118.
Carey, S.D. 1984. Amphiuma pholeter. Geographic distribution. Herpetological
Carey, S.D. 1985. Amphiuma pholeter. Geographic distribution. Herpetological
Conant, R., and J.T. Collins. 1998. Reptiles and Amphibians: Eastern/Central North
America. Houghton Miffl in Co., Boston, MA. 616 pp.
Daniels, R.J.R. 1992. Geographical distribution patterns in the Western Ghats, India.
Journal of Biogeography 19:521–529.
Dodd, C.K., Jr. 2004. The Amphibians of Great Smoky Mountains National Park.
The University of Tennessee Press, Knoxville, TN. 284 pp.
2010 S.P. Graham, D.A. Steen, K.T. Nelson, A.M. Durso, and J.C. Maerz 29
Gibbons, J.W., and K.A. Buhlmann. 2001. Reptiles and amphibians. Pp. 372–390,
In J.G. Dickson, (Ed.). Wildlife of Southern Forests: Habitat and Management.
Hanckock House Publishers, Blaine, WA. 552 pp.
Gibbons, J.W., Burke, V.J., J.E. Lovich, R.D. Semlitch, T.D. Tuberville, J.R. Brodie,
J.L. Greene, P.H. Niewiarowski, H.H. Whiteman, D.E. Scott, J.H.K. Pecnmann,
C.R. Harrison, S.H. Bennett, J.R. Krenz, M.S. Mills, K.A. Buhlmann, J.R. Lee,
R.A. Seigel, A.D. Tucker, T.M. Mills, T. Lamb, M.E. Dorcas, J.D. Congdon,
M.H. Smith, D.H. Nelson, M. B. Dietsch, H.G. Hanlin, J.A. Ott, and D.J. Karapatakis.
1997. Perceptions of species abundance, distribution, and diversity:
Lessons from four decades of sampling on a government-managed reserve. Environmental
Gotelli, N.J., and R.K. Colwell. 2001. Quantifying biodiversity: Procedures and
pitfalls in the measurement and comparison of species richness. Ecology Letters
Graham, S.P. 2008. Amphiuma pholeter. Geographic distribution. Herpetological
Graham, S.P. 2009. Desmognathus aeneus (Seepage Salamander). Geographic distribution.
Herpetological Review 40:232–233.
Graham, S.P., V.M. Johnson, and E.K. Timpe. 2007a. A bioblitz competition to assess
a distribution gap in Georgia herpetofaunal records. Herpetological Review
Graham, S.P., E.K. Timpe, and L.A. Giovanetto. 2007b. Significant new records for
Georgia herpetofauna. Herpetological Review 38:494–495.
Graham, S.P. V.M. Johnson, S.K. Hoss, D.A. Steen, D.J. McMoran, A.M. Durso, K.T.
Nelson, S.C. Cox, E.C. Susko, J.W. Shapiro, and J.R. McMillan. In press. An
additional bioblitz competition in northwest Georgia yields new county records.
Grenyer, R., C.D., Jackson, S.F., Thomas, G.H., Davies, R.G., Davies, T.J.,
Jones, K.E., Olson, V.A., Ridgly, R.S., Rassmussen, P.C., Ding, T.S., Bennett,
P.M., Blackburn, T.M., Gaston, K.J., Gittleman, J.L., and Owens, I.P. 2006.
Global distribution and conservation of rare and threatened vertebrates. Nature
Griffith, G.E., J.M. Omernik, J.A. Comstock, S. Lawrence, G. Martin, A. Goddard,
V.J. Hulcher, and T. Foster. 2001. Ecoregions of Georgia and Alabama (color
poster with map, descriptive text, summary tables, and photographs). US Geological
Survey, Reston, Virginia. (Map scale = 1:1,700,000).
Guyer, C., M. Bailey, J. Holmes, J. Stiles, and S. Stiles. 2007. Herpetofaunal response
to Longleaf Pine ecosystem restoration, Conecuh National Forest, Alabama.
Unpublished report to the US Forest Service, Auburn, University, Auburn,
AL. 92 pp.
Heyer, W.R., M.A. Donnelly, R.W. McDiarmid, L.-A.C. Hayek, and M.S. Foster
(Eds.). 1994. Measuring and Monitoring Biological Diversity: Standard Methods
for Amphibians. Smithsonian Institution Press, Washington, DC. 364 pp.
Jensen, J.B. 2000. Rana capito (Gopher Frog) geographic distribution. Herpetological
Jensen, J.B., and N.A. Klaus. 2004. Notophthalmus perstriatus (Striped Newt) Geographic
distribution. Herpetological Review 35:403.
Jensen, J.B., C.D. Camp, W. Gibbons, and M.J. Elliot. 2008. Amphibians and Reptiles
of Georgia. University of Georgia Press, Athens, GA. 600 pp.
30 Southeastern Naturalist Vol. 9, No. 1
Jones, S.B. 1974. The fl ora and phytogeography of the Pine Mountain region of
Georgia. Castanea 39:113–149.
Kerr, J.T., and L. Packer. 1997. Habitat heterogeneity as a determinant of mammal
species richness in high-energy regions. Nature 385:252–254.
Klaus, N.A., and J.B. Jensen. 2009. Micrurus fulvius (Harlequin Coralsnake). Geographic
distribution. Herpetological Review 40:365.
Luhring, T.M. 2008. “Problem species” of the Savannah River Site such as Brimley’s
Chorus Frog, Pseudacris brimleyi, demonstrate the hidden biodiversity
concept on an intensively studied government reserve. Southeastern Naturalist
Means, D.B. 1976. Survey of the status of amphibians and reptiles of the Apalachicola
National Forest, Florida. Unpublished report to the US Forest Service,
Tallahassee, FL. 60 pp.
Mount, R.H. 1975. The Reptiles and Amphibians of Alabama. Auburn University
Agricultural Experiment Station, Auburn, AL. 347 pp.
Myers, N. 1988. Threatened biotas: “Hotspots” in tropical forests. Environmentalist
Myers, N. 1990. The biodiversity challenge: Expanded hotspots analysis. Environmentalist
Myers, N., R.A. Mittermeier, C.G. Mittermeier, G.A.B. da Fonseca, and J. Kent.
2000. Biodiversity hotspots for conservation priorities. Nature 403:853–858.
National Park Service. 2009. Available online at http://nps.gov. Accessed 16 April 2009.
Palmer, W.M. 1987. A new species of glass lizard (Anguidae: Ophisaurus) from the
Southeastern United States. Herpetologica 43:415–423.
Patrick, T.S., J.R. Allison, and G.A. Krakow. 1995. Protected plants of Georgia.
Georgia Department of Natural Resources, Social Circle, GA.
Pianka, E.R. 1966. Latitudinal gradients in species diversity: A review of concepts.
American Naturalist 100:33–46.
Printiss, D., and D. Hipes. 1999. Rare amphibian and reptile survey of Eglin Air
Force Base, Florida. Unpublished Final Report to the Florida Natural Areas Inventory,
Tallahassee, FL. 65 pp.
Qian, H., and R.E. Ricklefs. 1999. A comparison of the taxonomic richness of vascular
plants in China and the United States. American Naturalist. 154:160–181.
Qian, H., X. Wang, S. Wang, and Y. Li. 2007. Environmental determinants of amphibian
and reptile species richness in China. Ecography 30:471–482.
Ricketts, T.H., E. Dinerstein, D.M. Olson, and C. Loucks. 1999. Who’s where in
North America? Patterns of species richness and the utility of indicator taxa for
conservation. BioScience 49: 69–380.
Rodrí guez, M.Á ., J.A. Belmontes, and B.A. Hawkins. 2005. Energy, water, and
large-scale patterns of reptile and amphibian species richness in Europe. Acta
Smith, L.L., D.A. Steen, J.M. Stober, M.C. Freeman, S.W. Golladay, L.M. Conner,
and J. Cochrane. 2006. The vertebrate fauna of Ichauway, Baker County, GA.
Southeastern Naturalist 5:599–620.
Steltenpohl, M.G., P.M. Mueller, A.J. Heatherington, T.B. Hanley, and J.L. Wooden.
2008. Gondwanan/peri-Gondwanan origin for the Uchee terrane, Alabama and
Georgia: Carolina zone or Suwannee terrane (?) and its suture with Grenvillian
basement of the Pine Mountain window. Geosphere 4:131–144.
Stevenson, D.J. 1999. The herpetofauna of Fort Stewart, Georgia: Habitat occurrence,
status of protected and rare species, and species diversity. Upublished
report to the Fort Stewart Fish and Wildlife Branch, Fort Stewart, GA. 98 pp.
2010 S.P. Graham, D.A. Steen, K.T. Nelson, A.M. Durso, and J.C. Maerz 31
Tilley, S.G., and J.E. Huheey. 2001. Reptiles and Amphibians of the Smokies. Great
Smoky Mountains Natural History Association, Gatlinburg, TN. 143 pp.
Todd, B.D., C.T. Winne, J.D. Willson, and J. W. 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
Tuberville, T.D., J.D. Willson, M.E. Dorcas, and J.W. Gibbons. 2005. Herpetofaunal
species richness of southeastern national parks. Southeastern Naturalist
US Fish and Wildlife Service (USFWS). 1998a. Okefenokee National Wildlife
Refuge amphibians, fish, mammals, and reptiles list. Unpublished pamphlet.
Folkston, GA. 16 pp.
USFWS. 1998b. St. Marks National Wildlife Refuge reptile, amphibian, and mammal
list. Unpublished pamphlet. St. Marks, FL. 8 pp.
Wharton, C.H. 1978. The Natural Environments of Georgia: Georgia Geologic Survey
Bulletin 114. 227 pp.
Zug, G.R. 1993. Herpetology. Academic Press. San Diego, CA. 527 pp.
32 Southeastern Naturalist Vol. 9, No. 1
Appendix 1. Amphibian and reptile species documented during a seven day survey in west-central Georgia in 2008, including new county records, locality data,
and voucher information. AU = Auburn University, UGA = University of Georgia. T = team responsible.
Species New record Locality Date Verified by T Museum number
Bufo terrestris Bonnaterre
Rana catesbeiana Shaw
Rana sphenocephala Webster Co. 32.051804°N, 84.548490°W 8-Oct-08 C. Guyer AU AHAP-D 135
R. clamitans Latreille
R. heckscheri Wright
R. capito LeConte
Hyla cinerea Schneider Webster Co. 32.051804°N, 84.548490°W 8-Oct-08 C. Guyer AU AHAP-D 134
Stewart Co. 32.141480°N, 84.753142°W 21-Mar-08 C. Guyer AU AHAP-D 113
H. squirella Bosc
H. gratiosa LeConte Webster Co. 32.107753°N, 84.492906°W 8-Oct-08 C. Guyer AU AHAP-D 133
H. avivoca Viosca Stewart Co. 32.034373°N, 84.890252°W 22-Mar-08 C. Guyer AU AHAP-D 120
Pseudacris crucifer Wied-Neuwied
P. feriarum Baird
P. ornata Holbrook
Acris crepitans Baird
A. gryllus LeConte
Scaphiopus holbrookii Harlan
Gastrophryne carolinensis Holbrook Webster Co. 32.051804°N, 84.548490°W 8-Oct-08 C. Guyer AU AHAP-D 136
Eurycea cirrigera Green
E. guttolineata Holbrook Talbot Co. 32.795118°N, 84.398492°W 10-Oct-08 C. Guyer AU AHAP-D 125
E. chamberlaini Harrison & Guttman Stewart Co. 32.034373°N, 84.890252°W 21-Mar-08 C. Guyer AU AUM 37748
Marion Co. 32.526694°N, 84.569986°W 22-Mar-08 C. Guyer AU AUM 37751
Plethodon glutinosus Green
P. websteri Highton
Desmognathus apalachicolae Means & Karlin
2010 S.P. Graham, D.A. Steen, K.T. Nelson, A.M. Durso, and J.C. Maerz 33
Species New record Locality Date Verified by T Museum number
D. conanti Rossman
Pseudotriton ruber Latreille
Gyrinophilus porphyriticus Green Talbot Co. 32.854411°N, 84.546829°W 26-Oct-08 C. Camp, J. Jensen AU AHAP-D 195-196
Ambystoma opacum Gravenhorst Schley Co. 32.308811°N, 84.293997°W 11-Oct-08 C. Guyer AU AHAP-D 126
A. talpoideum Holbrook Stewart Co. 32.139461°N, 85.041513°W 22-Mar-08 J.C. Godwin AU AUM 37759
A. maculatum Shaw Talbot Co. 32.795118°N, 84.398492°W 23-Mar-08 C. Guyer AU AHAP-D 117-118
Notophthalmus viridescens Rafinesque Webster Co. 32.051804°N, 84.548490°W 8-Oct-08 C. Guyer AU AHAP-D 137
Siren lacertina Linnaeus Schley Co. 32.174857°N, 84.371463°W 23-Mar-08 C. Guyer AU AUM 37760
Chelydra serpentina Linnaeus Talbot Co. 32.622125°N, 84.634668°W 23-Mar-08 C. Guyer AU AHAP-D 116
Macrochelys temminckii Harlan
Sternotherus minor Agassiz Schley Co. 32.174857°N, 84.371463°W 21-Mar-08 C. Guyer AU AHAP-D 194
S. odoratus Latreille
Kinosternon subrubrum Lacé pè de
Trachemys scripta Schoepff
Chrysemys picta Schneider
Pseudemys concinna LeConte
Deirochelys reticularia Latreille
Gopherus polyphemus Daudin
Terrapene carolina Schley Co. 32.215076°N, 84.371364°W 10-Oct-08 C. Guyer AU AHAP-D 122-123
Anolis carolinensis Voigt Schley Co. 32.174857°N, 84.371463°W 21-Mar-08 C. Guyer AU/UGA AHAP-D 108
Talbot Co. 32.786769°N, 84.422183°W 23-Mar-08 C. Guyer AU AHAP-D 200
Sceloporus undulatus Bosc & Daudin Schley Co. 32.174857°N, 84.371463°W 21-Mar-08 C. Guyer AU AHAP-D 192
Eumeces fasciatus Linnaeus Stewart Co. 32.181526°N, 84.824247°W 21-Mar-08 C. Guyer AU AHAP-D 110-112
E. laticeps Schneider Talbot Co. 32.799730°N, 84.400077°W 22-Mar-08 C. Guyer AU AUM 37761
Scincella lateralis Say Schley Co. 32.174857°N, 84.371463°W 21-Mar-08 C. Guyer AU AHAP-D 115
Elaphe obsoleta Holbrook
Coluber constrictor Linneaus Talbot Co. 32.799730°N, 84.400077°W 23-Mar-08 C. Guyer AU/UGA AHAP-D 119
34 Southeastern Naturalist Vol. 9, No. 1
Species New record Locality Date Verified by T Museum number
Masticophis fl agellum Shaw Schley Co. 32.206008°N, 84.317817°W 10-Oct-08 C. Guyer AU AHAP-D 121
Opheodrys aestivus Linnaeus Talbot Co. 32.750672°N, 84.419732°W 10-Oct-08 C. Guyer AU AHAP-D 124
Cemophora coccinea Blumenbach
Tantilla coronata Baird & Girard Schley Co. 32.308811°N, 84.293997°W 21-Mar-08 C. Guyer AU AUM 37758
Nerodia sipedon Linnaeus Talbot Co. 32.782546°N, 84.405132°W 22-Mar-08 C. Guyer UGA AHAP-D 201-202
N. erythrogaster Forster Schley Co. 32.174857°N, 84.371463°W 10-Oct-08 C. Guyer AU AHAP-D 127-128
Storeria dekayi Talbot Co. 32.782546°N, 84.405132°W 21-Mar-08 C. Guyer AU AHAP-D
Virginia valeriae Baird & Girard
Thamnophis sirtalis Linnaeus Webster Co 32.055479°N, 84.546561°W 8-Oct-08 C. Guyer AU AHAP-D 138
Diadophis punctatus Linnaeus Stewart Co. 32.181526°N, 84.824247°W 21-Mar-08 C. Guyer AU AHAP-D 109
Talbot Co. 32.799477°N, 84.502491°W 22-Mar-08 C. Guyer AHAP-D 114
Agkistrodon contortrix Linnaeus Stewart Co. 32.054132°N, 84.735728°W 8-Oct-08 C. Guyer AU AHAP-D 139
A. piscivorus Lacé pè de Schley Co. 32.174857°N, 84.371463°W 21-Mar-08 C. Guyer AU AHAP-D 193
Crotalus horridus Linnaeus
C. adamanteus Palisot de Beauvois