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22001177 SOUTHEASTERN NATURALIST 1V6o(2l.) :1266,1 N–2o8. 22
The Herpetofauna of Conservation Lands along the
Altamaha River, Georgia
Dirk J. Stevenson1,* and Houston C. Chandler1
Abstract - The Altamaha River region of southeastern Georgia is widely recognized for its
biodiversity and conservation value for imperiled species. Even so, intensive amphibian
and reptile surveys have never been conducted in this area. From 2008 to 2016, we conducted
herpetofaunal surveys at 13 conservation lands located along the Altamaha River and
along the lower reaches of a main tributary, the Ocmulgee River. We used a variety of field
methods including visual encounter surveys, turtle trapping, frog-call surveys, drift fences,
and surveys for snakes at Gopherus polyphemus (Gopher Tortoise) burrows. From these
data, combined with a review of the published literature and a search of relevant museum
collections, we determined that conservation and other nearby lands along the Altamaha
River support a diverse herpetofauna of 100 species, including 41 amphibian species (18
salamander and 23 frog and toad species) and 59 reptile species (1 crocodilian, 11 lizard, 33
snake, and 14 turtle species). Seventeen species (12 reptile and 5 amphibian species) that are
either federally listed, state-listed, or tracked as special concern by the Georgia Department
of Natural Resources have been documented here, and sandhills along the Altamaha River
support significant populations of Gopher Tortoises, Drymarchon couperi (Eastern Indigo
Snake), and Crotalus adamanteus (Eastern Diamond-backed Rattlesnake). Biogeographically,
the Altamaha River is a notable influence on the distributions of many amphibians and
reptiles. High species richness, including many declining species, underscores the Altamaha
River’s importance to conservation, and future efforts should focus on long-term monitoring
of imperiled species and effective management of conservation lands.
Introduction
Species lists may be developed through long-term accumulation of records or by
intense general collecting over a relatively short period, with long periods (usually
in the scales of years) needed to sample complex, diverse communities (Heyer et al.
1994). Developing site-specific lists of species is an important first step to conducting
long-term monitoring and/or research on specific populations and communities
(Tuberville et al. 2005). Significantly, knowledge of species present at a particular
site can assist in directing conservation efforts and habitat management goals. Over
long periods of time, with consistent effort, species lists can be used as a simple
means of monitoring plant and animal communities by identifying general patterns
in species occurrence without the intensive effort required for other monitoring
techniques (Droege et al. 1998, Roberts et al. 2007).
The Altamaha River, formed by the confluence of the Oconee and Ocmulgee
Rivers, flows 220 km through the Coastal Plain of southeastern Georgia before
1The Orianne Society, 11 Old Fruitstand Lane, Tiger, GA 30576. *Corresponding author -
dstevenson@oriannesociety.org.
Manuscript Editor: Brad Glorioso
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emptying into the Atlantic Ocean near Darien, GA. The Altamaha is the largest freeflowing
river on the east coast of the United States, with the lower Ocmulgee and
the entire Altamaha River main stem flowing unimpounded for a distance of about
480 km (Reese and Baxter 2007). The Altamaha watershed drains a huge portion
of Georgia (>3.6 million ha, 23% of the state), and the river discharges 378,000
L of fresh water into the Atlantic Ocean every second (Wharton 1978). A forested
swampy floodplain, up to 8 km wide and encompassing an estimated 68,800 ha,
buffers the Altamaha River main stem (Edwards et al. 2013).
An inventory of the Altamaha River watershed conducted by The Nature Conservancy
(TNC) in the 1990s documented 120 rare or endangered plant and animal
species (Dallmeyer and Ray 2012). Natural communities in the river floodplain
include some of the best remaining examples of bottomland hardwood forests
and cypress–tupelo swamps in Georgia (Edwards et al. 2013, Wharton 1978). In
recognition of its beauty, uniqueness, biodiversity, and conservation value, TNC
placed the Altamaha River on their list of “The 75 Last Great Places” (Edwards et
al. 2013). Within the last several decades, more than 40,000 ha along the Altamaha
River have been purchased and protected as conservation lands (Dallmeyer and Ray
2012, Edwards et al. 2013).
Although the Altamaha River region is known to support substantial biodiversity,
very little has been published specific to its herpetofauna, other than partial
species lists or distribution records (Schlimm 2013, Wharton 1978, Williamson
and Moulis 1994). To address this need, we conducted field inventories at a series
of protected conservation lands located along the lower Altamaha River (i.e., the
main stem Altamaha and the last ~30 km of the Ocmulgee). The goals of our study
were to provide contemporary species lists for these sites while contributing locality
records for populations of listed and imperiled species. Additionally, given the
overall size of our study area and the fact that the Altamaha River is of known biogeographic
importance to freshwater and terrestrial taxa (Liu et al. 2006, Page and
Burr 2011), we anticipated that our surveys would reveal biogeographic patterns or
distribution anomalies for some taxa.
Field-site Description
We conducted field surveys at 13 conservation sites (Wildlife Management
Areas [WMAs], natural areas, and private ecological preserves) located in the Altamaha
River region of southern Georgia (Table 1, Fig. 1). Study sites varied from
~1000 ha to ~12,000 ha (mean size = 4300 ha). The borders of 9 of these sites
follow the main stem of the Altamaha River. The other 4 sites are located in the
lower portion of the Ocmulgee River drainage and border 2 major tributaries to
the Altamaha River—the Ocmulgee River and the Little Ocmulgee River (Fig. 1).
In 2 cases, we surveyed conservation sites that were contiguous with each other
(Penholoway WMA and Sansavilla WMA; Orianne Indigo Snake Preserve and
Horse Creek WMA), and we combined amphibian/reptile occurrence data for
these sites.
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Table 1. Thirteen conservation sites in the Altamaha River region of southern Georgia that were surveyed from 2008 to 2016 for amphibian and reptile
species using a variety of standard herpetological sampling techniques (Foster 2012). Habitat types (following Edwards et al. 2013): 1 = sandhills and river
dunes, 2 = dry upland longleaf woodlands, 3 = dry evergreen oak woodlands, 4 = mesic slope forests/deciduous hardwood forests, 5 = pine flatwoods, 6
= small stream floodplain forests, 7 = seepage swamps/herb bogs/shrub bogs, 8 = cypress–gum ponds and depression marshes, 9 = bottomland hardwood
forests, 10 = cypress–tupelo river swamps, 11 = riverbanks and levees, 12 = tidal swamps, 13 = maritime hammocks. Survey methods: visual encounter
surveys (VS), drift fences with pitfall/funnel traps (DF), turtle trapping (hoop nets) (TH), turtle trapping (crab-wire Traps) (TC), snake surveys at Gopher
Tortoise burrows (GS), river edge snorkel/wade surveys (RS), anuran-call surveys (AS), minnow traps/aquatic funnel traps (AT), dipnet surveys (DS),
road-cruising (RC), amphibian–reptile bioblitzes (BB). Surveys were conducted in a variety of habitats over multiple years.
Site Area (ha) Dominant habitat types Survey methods Georgia counties Survey dates
Alligator Creek WMAA 3457 1, 2, 3, 5, 6, 7, 8, 10, 11 VS, DF, TC, GS, RS, AS, DS, RC Wheeler 2008–2012,
2015
Altama Plantation WMA 1674 1, 2, 3, 4, 5, 6, 7, 10, 12, 13 VS, TC, GS, AS, DS Glynn 2008, 2016
Altamaha WMA 12,387 1, 3, 4, 5, 7, 9, 10, 11, 12, 13 VS, TC, GS, RS, AS, DS, RC McIntosh 2013–2016
Big Hammock WMA 2795 1, 3, 5, 8, 9, 10, 11 VS, GS, RS, AS, DS, RC, BB Tattnall 2008–2016
Bullard Creek WMA 3908 2, 4, 5, 9, 10, 11 VS, RS, AS, DS Appling, Jeff Davis 2008–2016
Flat Tub WMA 2546 2, 4, 5, 7, 9, 10, 11 VS, GS, RS, AS, DS Coffee, Jeff Davis 2008–2016
Griffin Ridge WMA 2756 1, 3, 5, 8, 9, 10, 11 VS, TC, GS, RS, AS, DS, RC, BB Long 2008–2016
Horse Creek WMA 3017 2, 4, 5, 8, 9, 10, 11 VS, TH, TC, GS, RS, AS, DS, BB Telfair 2008–2016
Moody Forest Natural Area 1801 2, 4, 5, 6, 7, 9, 10, 11 VS, TH, TC, GS, RS, AS, DS, RC, BB Appling 2008–2016
Orianne Indigo Snake Preserve 1023 1, 2, 4, 5, 7, 9, 10, 1 1 VS, DF, TH, TC, GS, RS, AS, AT, DS, BB Telfair 2008–2016
Penholoway Swamp WMA 4415 2, 4, 5, 6, 7, 8, 9, 10, 1 1 VS, GS, RS, AS, RC Wayne 2008–2016
Sansavilla WMA 6781 2, 4, 5, 6, 7, 8, 9, 10, 11 VS, GS, RS, AS, DS, RC Glynn, Wayne 2008–2016
Townsend WMA 11,648 1, 3, 4, 5, 7, 8, 9, 10, 11 VS, TC, GS, RS, AS, AT, DS, RC, BB Long, McIntosh 2008–2016
ASurveys at Alligator Creek WMA included ~2200 ha of well-managed land on private property adjacent to the WMA.
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Methods
From January 2008 through September 2016, we surveyed individual sites
on 5–20 dates annually; some sites were surveyed in fewer years (Table 1). We
used a wide variety of standard herpetological sampling techniques, including visual
encounter surveys (turning surface cover, inspecting logjams and brush piles,
checking rotten logs and snags, etc.), drift fences with pitfall and/or funnel traps,
turtle trapping with conventional hoop net traps, turtle trapping with modified crabwire
traps, snake surveys at Gopherus polyphemus (Gopher Tortoise) burrows,
wading surveys of river-edge habitats, anuran-call surveys, sampling wetlands
with aquatic minnow traps or funnel traps, dipnet surveys, and road-cruising (Table
1; see Heyer et al. 1994 and Foster 2012 for detailed descriptions of most of the
above methods). We used custom-made crab-wire turtle traps that measured approximately
61.0 cm x 61.0 cm x 47.0 cm and were made of 16-gauge galvanized
steel-mesh wire coated with black polyvinyl chloride. These traps are effective in
capturing small emydid (e.g., Clemmys guttata [Spotted Turtle]) and kinosternid
turtles (Chandler et al., in press). We conducted cool-season snake surveys at tortoise
burrows, targeting Drymarchon couperi (Eastern Indigo Snake) and Crotalus
adamanteus (Eastern Diamond-backed Rattlesnake), by visiting sandhill habitats
supporting tortoise populations from November–March and searching for basking
snakes and shed skins (Bauder et al. 2017). In addition to the above survey methods,
we organized bioblitz events at some of our study sites, wherein participants
divided into teams and surveyed intensively over 2–3-day periods in an effort to
document as many amphibian and reptile taxa as possible (Table 1). In addition
Figure 1. Conservation sites along the Altamaha River, GA, and the lower Ocmulgee River,
GA, that were sampled for herpetofauna using a variety of techniques from 2008 to 2016.
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to surveys of the 13 primary survey sites, we conducted visual encounter surveys
targeting a semi-aquatic viper, Agkistrodon piscivorus (Cottonmouth), at floodplain
swamps located along the entire length of the Altamaha River.
We attempted to survey all representative habitats present at each study site
(Table 1). We placed an emphasis on the following habitats: (1) the main stem
of the major river systems that served as study site boundaries (Altamaha, Little
Ocmulgee, and Ocmulgee rivers); (2) habitats within the floodplains of these river
systems, including bottomland hardwood forests, Taxodium distichum (L.) Rich.
(Bald Cypress)–Nyssa aquatica L. (Water Tupelo) swamps and sloughs, and oxbow
lakes; (3) perennial, mucky springs and seepage areas located at the base of sandhills
or in ravines associated with north-facing bluffs; (4) isolated, depressional
wetlands (e.g., Taxodium ascendens Brongn. [Pond Cypress], Nyssa sylvatica
Marshall [Black Gum], and Ilex myrtifolia Walter [Myrtle Dahoon] wetlands) embedded
in pine uplands (pine flatwoods, sandhills); and (5) Pinus palustris Mill.
(Longleaf Pine)–Aristida stricta Michaux (Wiregrass) flatwoods and sandhills.
A notable weakness of wildlife surveys, and herpetological surveys in particular,
is that some species present on-site may elude detection because of low activity
levels, low abundance, and/or clandestine life histories (e.g., fossorial species)
(Mazerolle et al. 2007, O’Donnell and Semlitsch 2015). Thus, survey effort and
seasonality of surveys has a profound effect on survey completeness, especially
with regard to cryptic species that occur at low densities or are distributed in habitats
that are hard to sample (Foster 2012). Considering this, we conducted surveys
year-round and scheduled surveys such that they coincided with individual species’
phenology and with weather and soil-moisture conditions optimal for amphibian
and reptile activity.
We developed a hypothetical species list for the Altamaha River region (Appendix
1) by consulting range maps in current herpetological references (e.g.,
Jensen et al. 2008, Powell et al. 2016). We did not include Malaclemys terrapin
(Schoepff) (Diamond-backed Terrapin), a coastal species, or sea turtle species on
this list. With respect to taxonomy, we follow Crother (2012), except that we recognize
Pseudemys floridana (Coastal Plain Cooter) as a full species. A number of
other herpetologists currently recognize this form as a full species (e.g., Guyer et
al. 2015, Jensen et al. 2008, Powell et al. 2016), and we do so because this species
is morphologically distinguishable from Pseudemys concinna (River Cooter) in our
study area.
To locate additional and historic herpetofaunal records for our study sites, we
reviewed the published literature, conducted a search of relevant museum collections,
and interviewed local herpetologists and natural resource managers (with
the Georgia Department of Natural Resources, TNC, and the US Fish and Wildlife
Service) who had personal field experience with some or all of our study sites.
To locate literature records, we consulted Jensen et al. (2008), Schlimm (2013),
and Williamson and Moulis (1994). To locate museum records, we searched the
following museum collections: Auburn University (AUM), Florida State University
(FSU), Georgia Museum of Natural History (GMNH), Georgia Southern
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University (GSU), Georgia Southwestern State University (GSWSU), Florida
Museum of Natural History (FMNH), and Valdosta State University (VSU). Other
major museum collections were searched as recently as 1994 by Williamson and
Moulis (1994). To document a species occurrence at a particular site, for specimens
representing range extensions or county records and for records of rare or listed
taxa, we deposited voucher specimens (or photographs) in museum collections
including the Georgia Museum of Natural History (GMNH), Georgia Southern
University (GSU), the Florida Museum of Natural History (FMNH), and the North
Carolina State Museum of Natural Sciences (NCSM) (see Supplementary File 1,
available online at http://www.eaglehill.us/SENAonline/suppl-files/s16-2-S2350-
Stevenson-s1, and, for BioOne subscribers, at http://dx.doi.org/10.1656/S2350.s1).
Results and Discussion
Species richness
Our hypothetical amphibian and reptile species list for the Altamaha River
region of southern Georgia totaled 106 species and was comprised of 45 amphibian
species (21 salamander and 24 frog and toad species) and 61 reptile species
(1 crocodilian, 14 turtle, 12 lizard, and 34 snake species) (Appendix 1). Overall, for
the 13 conservation land sites that we surveyed as part of this study, we documented
(including museum records) 36 amphibian species (13 salamander and 23 frog and
toad species) and 57 reptile species (1 crocodilian, 14 turtle, 10 lizard, and 32 snake
species) (Appendix 1). From our field work in other parts of the Altamaha River
region (outside of the 13 conservation land sites) and from museum specimens,
we located records for an additional 5 salamander species, 1 lizard species, and 1
snake species within 5 km of the Altamaha River main stem or its floodplain (Appendix
1). We failed to document, or locate museum records for, 4 amphibian and 2
reptile species from our hypothetical species list. No introduced species were found
at any of our survey sites.
The total number of species we documented from conservation lands varied
from 49 to 75, with the greatest species richness recorded at Orianne Indigo Snake
Preserve/Horse Creek WMA (75), Moody Forest Natural Area (71), and Big Hammock
WMA (70) (Fig. 2). We documented 55 new county records and several range
extensions during this study (e.g., Butler et al. 2012; Jensen et al. 2011; Stevenson
et al. 2009a, 2011). Our study demonstrates that the species richness for the Altamaha
River region is high, both overall and at individual conservation sites. Other
Coastal Plain sites in Georgia known to support comparable or higher diversity of
amphibians and reptiles include the Okefenokee National Wildlife Refuge (88 species;
Smith et al. 2006a; L. Smith, Joseph W. Jones Ecological Research Center at
Ichauway, pers. comm.); Fort Stewart Military Installation (97 species; Stevenson
1999); and Joseph W. Jones Ecological Research Center at Ichauway (86 species,
Smith et al. 2006b; L. Smith, Joseph W. Jones Ecological Research Center at Ichauway,
pers. comm.). We suspect that additional field work, particularly surveys using
drift fences, would document additional amphibian and reptile species from some
of the 13 conservation lands. The results of our inventory may have overlooked
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difficult-to-find species like fossorial snakes and salamanders, glass lizards, and
other clandestine species.
During our 2008–2016 surveys, we recorded all 14 turtle species expected
from the region, including 12 freshwater turtle species. We documented River Cooter
populations in the main stems of the lower Oconee and lower Ocmulgee rivers, and
throughout the Altamaha River main stem as far downstream as Fort Barrington, Mc-
Intosh/Wayne counties (Stevenson et al. 2012). Previously, this turtle was unreported
from the Altamaha River proper (Ward and Jackson 2008). We documented Sternotherus
minor (Loggerhead Musk Turtle) at 6 of 11 (54%) survey sites and found
this species elsewhere in the Altamaha River main stem as far downstream as Jesup,
Wayne/Long counties, where the species is currently abundant (D.J. Stevenson,
unpubl. data). Downstream of Jesup, we found only a single example of this molluskeating
turtle, at Townsend WMA in Long County. We suspect the Loggerhead Musk
Turtle population may have declined (downstream of Jesup) in association with declines
of native pearly mussels (Unionidae) from this section of the Altamaha River
(Meador 2008; J. Wisniewski, Georgia DNR, Social Circle, GA, pers. comm.). Although
the Altamaha River supports large turtle populations, a diverse turtle fauna,
and is located within a southeastern United States Turtle Priority Area (Buhlmann
et al. 2009), its freshwater turtle fauna has never been intensively studied. We recommend
the initiation of long-term monitoring programs at select freshwater sites
throughout the Altamaha drainage.
Figure 2. Species richness at 13 conservation sites along the Altamaha River, GA, that were
surveyed from 2008 to 2016. Herpetofaunal observations were pooled for Orianne Indigo
Snake Preserve (OISP)/Horse Creek WMA and for Penholoway Swamp/Sansavilla WMA
because these sites share a border with each other. The “total species” bar represents the
maximum number of species that potentially occur in this region.
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Despite targeting seepage habitats, we were only able to document Pseudotriton
ruber (Red Salamander) at 2 of 11 (18%) survey sites. In the Coastal Plain, this species
inhabits perennial, mucky spring-seepage habitats located at the base of xeric
sandhills or in ravines associated with north-facing bluffs (Means 2000, Stevenson
et al. 2009c). An Altamaha River bluff (located near Jesup, Wayne County; not one
of our survey sites) known to harbor a population of Red Salamanders is the southeastern-
most site known for this species (Williamson and Moulis 1994). Perennial
seepages, locally distributed environments in the Coastal Plain, typically support
a high diversity and biomass of salamanders (Means 2000, Stevenson et al. 2009c),
and we encourage natural resource managers in the Altamaha River region to actively
identify and safegaurd these habitats where they occur on protected lands.
Intriguingly, we recorded Cottonmouths from only 5 of 11 (45%) survey sites,
and we were unable to document this species from alluvial river swamps along
most of the length of the Altamaha River main stem, despite additional survey effort.
In the Altamaha River floodplain, except for 2 sites (both significantly altered
wetlands) in Long County, we routinely observed Cottonmouths only from the
southeastern-most stretches of the Altamaha River system in Glynn and McIntosh
counties (Fig. 3). Here, we found (or compiled records for) Cottonmouths from
river swamps, tidal marshes, and other wetland habitats (including on Chapney,
Butler, and Lewis islands) from about river-km 48 east to the coast. The only
credible Cottonmouth records obtained by our interviews were from this area. In
Figure 3. Agkistrodon piscivorus (Cottonmouth) observations and museum records for
the Altamaha River region of southeastern Georgia. Survey sites were sampled for Cottonmouths
on a minimum of 10 dates from 2008 to 2016. Study observations outside of
the Altamaha River floodplain were made by the authors from 2000 to 2016 via chance
encounters.
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contrast, we documented 1–3 species of watersnakes (Nerodia spp.) in river swamp
habitats at all of our survey sites and throughout the region (Appendix 1; Stevenson
and Stackhouse 2012).
In the Coastal Plain of southeastern Georgia (including sites proximal the Altamaha
River) and northern Florida, Cottonmouths have been recorded from coastal
marshes, blackwater creek swamps, isolated depressional wetlands (cypress ponds,
gum ponds, Carolina bays) embedded in mesic pine flatwoods, and grassy roadside
ditches, among other wetland habitats (Erwin et al. 2016; Gibbons and Semlitsch
1991; Williamson and Moulis 1979, 1994; Wolfe et al. 1988). Dorcas and Gibbons
(2008) mention that Cottonmouths are very common in many areas of Georgia and
describe them as the “classic venomous snake of southern river swamps”. Cottonmouths
are prone to utilize vegetated shorelines with abundant basking objects
and congregate around receding waters in swampy floodplains where prey is concentrated
(Palmer and Braswell 1995, Trauth et al. 2004), and we regularly focused
our surveys on these microhabitats when searching for Cottonmouths at Altamaha
River floodplain sites.
We believe that a recent Altamaha River drainage-wide decline of Cottonmouth
populations is unlikely, and, considering the intensity and duration of our survey
efforts, we feel it is unlikely that we overlooked the species. Instead, we conclude
that Cottonmouths are indeed absent (or extraordinarily rare) from much of the
Altamaha River and its floodplain. We posit that a perpetual instability (i.e., dramatically
fluctuating water levels driven by large annual pulses in flow; frequent
flooding and then drying) of aquatic habitats in the Altamaha River floodplain may
be responsible, at least in part, for the absence of Cottonmouths. Regular scouring
of the floodplain may remove graminaceous vegetation (i.e., snake cover) from
the margins of sloughs in the floodplain and/or influence distribution of prey. The
presence of moving water in the floodplain may in some way deter Cottonmouth
presence (Wharton 1978). The apparent lack of Cottonmouth populations at many
floodplain swamps along the Altamaha River merits further study.
Imperiled species
Our surveys of Altamaha River conservation lands documented the federally
listed Eastern Indigo Snake and 4 other species that are state-listed in Georgia
(Lithobates capito [Gopher Frog], Gopher Tortoise, Spotted Turtle, and Heterodon
simus [Southern Hog-nosed Snake]) (Table 2). For example, we documented
the Spotted Turtle at 5 of 11 (45%) survey sites, generally inhabiting blackwater
creek swamps and seepage slope swamps. This species has never been documented
from alluvial river swamps along the lower Ocmulgee River or the Altamaha
River (Stevenson et al. 2015). Additionally, we recorded 1 amphibian (Desmognathus
auriculatus [Southern Dusky Salamander]) and 7 reptile species (2 lizard
[Ophisaurus attenuatus (Slender Glass Lizard), Plestiodon egregius (Mole
Skink)] and 5 snake species [Eastern Diamond-backed Rattlesnake, Farancia erytrogramma
(Rainbow Snake), Micrurus fulvius (Harlequin Coralsnake), Pituophis
melanoleucus (Pinesnake), Rhadinaea flavilata (Pine Woods Littersnake)]) that
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are tracked as species of special concern by the Georgia Department of Natural
Resources (Table 2).
We documented Eastern Indigo Snakes at 8 of 11 (73%) survey sites and Eastern
Diamond-backed Rattlesnakes (a declining, species of special concern) at 10 of 11
sites (91%). In south Georgia, adult Eastern Indigo Snakes are semi-migratory and
have very large home ranges that include xeric sandhills (where they overwinter in
Table 2. Amphibian and reptile species of conservation concern in the Altamaha River region of southern
Georgia (rare = RA, threatened = TH, candidate = CA, unusual = UN). Species that do not have
official state or federal status are listed as species of special concern (SOC) by the Georgia Department
of Natural Resources. Some species were documented on 1 or more of 13 conservation sites by
surveys conducted from 2008 to 2016, whereas others are known from historic records for this region.
Species with no historic or current records may occur or have occurred in this region as their ranges
overlap the region (Jensen et al. 2008, Powell et al. 2016).
# of conservation sites with Total # of
Federal State pre-2008 records from conservation
Species status status records 2008–2016 sites with records
Anurans
Lithobates capito RA 1 1 2
Lithobates virgatipesA SOC 0
Salamanders
Ambystoma cingulatumA TH TH 0
Ambystoma tigrinumB SOC 0
Notophthalmus perstriatusA CA TH 0
Desmognathus auriculatus SOC 1 1
Stereochilus marginatusB SOC 0
Necturus punctatusA SOC 0
Pseudobranchus striatusB SOC 0
Lizards
Ophisaurus attenuatus SOC 1 1 2
Ophisaurus compressusB SOC 0
Ophisaurus mimicusA RA 0
Plestiodon egregius SOC 2 2
Snakes
Crotalus adamanteus SOC 2 11 11
Drymarchon couperi TH TH 4 9 9
Farancia erytrogramma SOC 1 5 6
Heterodon simus TH 1 1 1
Micrurus fulvius SOC 1 7 7
Pituophis melanoleucus SOC 5 5
Rhadinaea flavilata SOC 1 1
Seminatrix pygaeaA SOC 0
Turtles
Clemmys guttata UN 5 5
Gopherus polyphemus CA TH 5 11 11
ASpecies that occur on the hypothetical species list for the region but have not been documented anywhere
within 5 km of the Altamaha River main stem.
BSpecies that were not documented from the 13 conservation land sites but are known from within 5
km of the Altamaha River main stem based on museum records.
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Gopher Tortoise burrows [Diemer and Speake 1983; Stevenson et al. 2003, 2009b])
as well as lower, wetter habitats (pine flatwoods, swamps) used by foraging snakes
during the warmer months (Hyslop et al. 2014). Longleaf Pine–Wiregrass sandhill
habitats located on xeric sand ridges and other well-drained soils along the Ocmulgee
and Altamaha Rivers support notably large Gopher Tortoise populations
(over 250 adults) present at 5 of 11 (45%) survey sites (M. Elliott, Georgia DNR,
Social Circle, GA, pers. comm.). The especially large home ranges of Eastern Indigo
Snakes (~1400 ha for some adult males in southern Georgia) have led experts to
classify this snake as an “umbrella species” (Hyslop 2007, Hyslop et al. 2014).
A number of other amphibian and reptile taxa endemic to Longleaf Pine habitats,
all of which have or are currently experiencing population declines (Jensen et
al. 2008, Means 2006), were not found by our surveys (e.g., Ophisaurus mimicus
[Mimic Glass Lizard], Notophthalmus perstriatus [Striped Newt], and Ambystoma
cingulatum [Frosted Flatwoods Salamander]) or were found at only 1–2 sites
(e.g., Gopher Frog and Southern Hog-nosed Snake). Many of the survey properties
either lacked isolated depressional wetlands (breeding habitat for the Gopher
Frog, Striped Newt, and Frosted Flatwoods Salamander), or depressional wetlands
on-site had been significantly and adversely impacted by drought, long-term fire
suppression, and, historically, commercial forestry activities. These impacts diminished
habitat suitability and may have led to the extirpation of these amphibian
species at these sites, if they were ever present (Bishop and Haas 2005, Chandler
et al. 2016, Means and Means 2005). We recommend that isolated and depressional
wetlands be a focus of restoration and management actions to improve the quality
of these often overlooked habitats.
Protecting large landscapes (≥2000 ha), with intact corridors and naturally functioning
uplands and wetlands will conserve Eastern Indigo Snakes while also providing
habitat for a broad diversity of other plant and animal species (Hyslop 2007).
At this time, the Ocmulgee–Altamaha region of southern Georgia is considered a
significant population stronghold for Gopher Tortoises, Eastern Indigo Snakes, and
Eastern Diamond-backed Rattlesnakes (M. Elliott, pers. comm.; Enge et al. 2013;
Stohlgren et al. 2015). We encourage continued monitoring of these species at these
conservation land sites and throughout the Altamaha region to detect population
trends through time (e.g., Bauder et al. 2017).
Despite intensive visual encounter survey efforts seeking the species, our
surveys did not document the Southern Dusky Salamander (a declining species
of special concern) from any of the Altamaha conservation lands. In southern
Georgia, this species and Stereochilus marginatus (Many-lined Salamander), also
not documented by our study, are most commonly associated with muck-floored,
acidic blackwater creek swamps, a wetland habitat different from the alluvial
swamps fringing the main stem of the Altamaha River (Graham et al. 2010, Means
2000, Stevenson and Stackhouse 2012). Blackwater creek swamp habitats were not
widespread on most of our survey sites, although we did locate potentially suitable
habitat on Altamaha WMA in McIntosh County, for which there is a historic museum
record for Southern Dusky Salamander. A specimen has recently been collected
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from a blackwater creek swamp that is a tributary of the Altamaha River, outside of
our survey area (Beamer and Lamb 2008, Graham et al. 2010).
Biogeography
The Altamaha River is a significant barrier to, or influence on, the distributions
of a number of amphibian and reptile species, as well as a number of other taxa
(e.g., fish [Page and Burr 2011], crayfish [Hobbs 1981], and the periodical cicada
Magicicada tredecim (Walsh and Riley) [Hinkle et al. 2012]). Our surveys and
review of museum records revealed that 5 salamander, 3 frog, 1 turtle, and 4 snake
species have ranges that, in the Atlantic Coastal Plain of Georgia, do not extend
south of the Altamaha River (and its associated floodplain) (Table 3). Some of these
species (e.g., Ambystoma tigrinum [Eastern Tiger Salamander] and Agkistrodon
contortrix [Copperhead]) are undocumented from the south side of the Altamaha
River main stem. Four of these amphibian species (Ambystoma maculatum [Spotted
Salamander], Desmognathus conanti [Spotted Dusky Salamander], Pseudacris
feriarum [Upland Chorus Frog], and Acris crepitans [Eastern Cricket Frog]) have
ranges that extend from the Piedmont into the Atlantic Coastal Plain of southeastern
Georgia, following the Altamaha River drainage (including the Oconee and
Table 3. Amphibian and reptile species whose ranges are influenced by the Altamaha River drainage
in southern Georgia.
Range extends into Altamaha R. is the In Georgia, species
Coastal Plain of southeastern-most is absent from
southeastern GA extent of range in the north of the Altamaha R.
Species along the Altamaha R. Atlantic Coastal Plain to the Savannah R.
Amphibians
Acris crepitans X X
Ambystoma tigrinumA X
Ambystoma maculatum X X
Desmognathus conanti X X
Hyla avivoca X
Necturus punctatus X
Pseudacris feriarum X X
Pseudotrition ruberB X
Reptiles
Agkistrodon contortrix X
Nerodia erythrogasterC X
Pseudemys concinna X
Rhadinaea flavilata X
Storeria dekayi X
AA 2015 A. tigrinum record for 3.3 km N of the main stem of the Altamaha River, Wheeler County,
GA (GMNH 50939) is one of the few for this taxa within the Altamaha River drainage.
BWe examined a P. ruber specimen (SSM 9474, now GSU 20029) from the Satilla River drainage
(Coffee Co., GA) catalogued by Williamson and Moulis (1994) and reclassified it as a P. montanus
(Mud Salamander).
CThere is a single literature record for this species south of the Altamaha River in the Satilla River
drainage (Neill 1946).
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2017 Vol. 16, No. 2
Ocmulgee Rivers) (Table 3). All of these amphibians occupy riparian corridors of
mesic bottomland forest that fringe (and are within the floodplains of) these river
systems, and we documented all of these species as far southeast (i.e., downstream)
along the Altamaha River as Baxley (Appling County) or Jesup (Wayne County),
but not beyond (Jensen et al. 2011; Stevenson et al. 2009a, 2011). The Appling/
Wayne counties region coincides with the inland-most portion of the Lower Coastal
Plain (located near Jesup), demarcated by the Wicomico Terrace, the oldest Pleistocene
shoreline (elevation ~30 m and ~1.5 million years old) (Gore and Witherspoon
2013, Wharton 1978).
There are several species whose ranges only partially overlap the Altamaha
River drainage. The Pine Woods Littersnake, a Coastal Plain species ranging from
North Carolina to Louisiana, including much of Florida, was documented from only
1 (9%) survey site. This species has never been documented anywhere in Georgia
north of the Altamaha River (i.e., between the Altamaha River and the Savannah
River) despite being known from adjacent South Carolina, and from southeastern
Georgia south of the Altamaha River. Intensive efforts, including on a public land
(Fort Stewart) north of the Altamaha that contains extensive suitable habitat (Stevenson
1999, Williamson and Moulis 1979), have been unsuccessful in finding this
species. The southernmost range extent of Necturus punctatus (Dwarf Waterdog) is
purported to be the Altamaha River drainage (Jensen et al. 2008). In actuality, the
species has never been verified from the main stem of the Altamaha River or from
any of its Lower Coastal Plain tributaries. Our efforts to record this salamander
were unsuccessful, and we suspect that water temperatures (too warm) may influence
this species’ southernmost distribution.
Conclusions
The Altamaha River region of Georgia supports an impressive herpetofaunal
assemblage, including many rare and declining species. We documented many species
at a suite of conservation lands throughout the region, and these sites support
some of the best remaining habitats in the Coastal Plain of southeastern Georgia.
Despite high species richness and habitat quality, few studies have focused on this
region of the state, and there is little or no long-term monitoring for most species.
We believe that this region of Georgia is of significant conservation value, and we
encourage additional habitat management/restoration and research to ensure that it
remains a hotspot for biodiversity.
Acknowledgments
We thank the museum curators and collection managers who provided us with records
and assisted in locating and depositing specimens, including L. McBrayer (Georgia Southern
University), N. Castleberry (University of Georgia), D. Laurencio and B. Folt (Auburn University),
E. Lemmon (Florida State University), K. Krysko (University of Florida Museum
of Natural History), and R. Moulis (Savannah Science Museum). The following people made
significant contributions to this study by assisting with field surveys and/or providing recent
observations: J. Bauder, D. Beamer, K. Briggs, A. Day, M. Elliott, M. Ishimatsu, J. Holland,
C. Howe, C. Jenkins, J. Jensen, B. Knight, J. Oguni, P. Moler, M. Moore, J. Palis, J. Parker, K.
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Ravenscroft, M. Ravenscroft, A. Safer, E. Schlimm, F. Snow, B. Stevenson, M. Stoddard, K.
Stohlgren, W. Vaigneur, and M. Wallace Sr. R. Horan provided numerous amphibian/reptile
records for the lower Altamaha region. M. Elliott, M. Hodges, J. Lee, and C. Martin helped
us access properties owned by the Georgia Department of Natural Resources and The Nature
Conservancy. We are especially grateful to the conservation-minded herpetological enthusiasts
and Orianne Society supporters who participated in organized amphibian and reptile
bioblitz events. Field work for this study was carried out under scientific collecting permits
from the Georgia Department of Natural Resources (29–WJH–16–21) and the US Fish and
Wildlife Service (TE28025A-2). We thank 2 anonymous reviewers and Brad Glorioso for
their helpful comments on an earlier draft of this manuscript.
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Appendix 1. Amphibian and reptile species known or with the potential to occur in the Altamaha River region of southern Georgia. Methods used to
document species: 1 = found by this survey, 2 = found by this survey; voucher specimen deposited in museum collection, 3 = literature record from
Schlimm (2013), 4 = personal communication to authors, 5 = museum specimen, collected pre-2000, 6 = museum specimen, collected post-2000. Abbreviations
for each area (and individuals who supplied information for that area through personal communications): AC = Alligator Creek WMA; AP
= Altama Plantation WMA (A. Day, M. Elliott, R. Horan, M. Stoddard); A = Altamaha WMA (R. Horan, M. Stoddard); BH = Big Hammock WMA (A.
Day, W. Vaigneur); BC = Bullard Creek WMA (M. Elliott); FT = Flat Tub WMA (M. Elliott, M. Stoddard); GR = Griffin Ridge WMA (D. Beamer, J.
Jensen); HC /OIS = Horse Creek WMA and Orianne Indigo Snake Preserve; MF = Moody Forest NA (J. Beane, C. Martin, P. Moler); PS/S = Penholoway
Swamp WMA and Sansavilla WMA (J. Evans); and T = Townsend WMA (M. Elliott, J. Jensen).
Wildlife Management Area, Natural Area, or Preserve
Species AC AP A BH BC FT GR HC/OIS MF PS/S T
Anurans
Anaxyrus terrestris (Bonnaterre) (Southern Toad) 1 1, 5 1, 5 1 1 1 1 1, 3 1, 3, 6 1 1, 3
Anaxyrus quercicus (Holbrook) (Oak Toad) 1 4 1 4 4 1 1, 3
Gastrophryne carolinensis (Holbrook) (Eastern 1 1, 5 1 1, 5 1 1 1 1, 3 1, 3 1 1, 3
Narrow-mouthed Toad)
Pseudacris crucifer (Wied-Neuwied) (Spring Peeper) 1 1 1 1 1 1 1, 5 1 1 1 1, 5
Pseudacris nigrita (Le Conte) (Southern Chorus Frog) 1 1, 6 1 1 1 1 1
Pseudacris feriarum (Baird) (Upland Chorus Frog) 1, 6 2 2 1, 6
Pseudacris ocularis (Bosc and Daudin) (Little Grass Frog) 1 1, 5 1 1 1 1 1, 5 1, 6 1 1 1
Pseudacris ornata (Holbrook) (Ornate Chorus Frog) 5 2
Acris crepitans Baird (Eastern Cricket Frog) 1 2 2 1 2 1, 6
Acris gryllus (Le Conte) (Southern Cricket Frog) 1 1, 5 1 1 1 1 1, 5 1, 3 1, 3 1 1, 5
Hyla avivoca Viosca (Bird-voiced Treefrog) 2 1 1 1 1 2 1 1
Hyla chrysoscelis Cope (Cope’s Gray Treefrog) 2 1 1 1 1 1 1 1 1
Hyla squirella Bosc (Squirrel Treefrog) 1 1 1 1 1 1 1 1 1 1 1
Hyla femoralis Bosc (Pine Woods Treefrog) 1 4 1 1 1 1 1, 3 1, 3 1 1, 3
Hyla gratiosa Le Conte (Barking Treefrog) 1 1 1, 3 3 1 1
Hyla cinerea (Schneider) (Green Treefrog) 1 1 1, 5 1 1 1 1 1 1, 3, 6 1 1
Scaphiopus holbrookii (Harlan) (Eastern Spadefoot) 2 1 1 1, 3 3 2, 3
Lithobates sphenocephalus (Cope) (Southern Leopard Frog) 1 1, 5 1, 5 1, 5 1 2 1 1, 3 1, 3, 6 1 1, 3
Lithobates capito (Le Conte) (Gopher Frog) 2 5
Lithobates catesbeianus (Shaw) (American Bullfrog) 1 1, 5 1 1 1, 5 1 1 1 1, 3 1 1
Lithobates clamitans (Latreille) (Green Frog) 1 1, 5 1 1 1 1, 5 2 1, 3, 6 1 1
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Wildlife Management Area, Natural Area, or Preserve
Species AC AP A BH BC FT GR HC/OIS MF PS/S T
Lithobates heckscheri (Wright) (River Frog) 1 1 2 1 1, 6 1
Lithobates grylio (Stejneger) (Pig Frog) 2, 5 1, 5 5
Lithobates virgatipes (Cope) (Carpenter Frog)A
Salamanders
Ambystoma cingulatum Cope (Frosted Flatwoods Salamander)A
Ambystoma opacum (Gravenhorst) (Marbled Salamander) 1 1 2 1 5 1, 3 1, 6
Ambystoma maculatum (Shaw) (Spotted Salamander) 1, 5 2 1 2
Ambystoma talpoideum (Holbrook) (Mole Salamander)B
Ambystoma tigrinum (Green) (Eastern Tiger Salamander)B
Amphiuma means Garden (Two-toed Amphiuma) 2 2 1 2 2 1 4 1 1
Notophthalmus perstriatus (Bishop) (Striped Newt)A
Notophthalmus viridescens (Rafinesque) (Eastern Newt) 2 1 1 2 1 1, 5 1 1, 6 2, 5
Desmognathus auriculatus (Holbrook) (Southern Dusky 5
Salamander)
Desmognathus conanti Rossman (Spotted Dusky Salamander)B
Eurycea cirrigera (Green) (Southern Two-lined Salamander) 1 1 1 1 4 1 1, 6 1 1
Eurycea guttolineata (Holbrook) (Three-lined Salamander) 1 2 5 1 1, 6 1
Eurycea quadridigitata (Holbrook) (Dwarf Salamander) 1 1 1 1, 5 1 1 1, 5 1 1, 6 1, 5 1
Plethodon ocmulgee Highton (Ocmulgee Slimy Salamander) 1 1 1 1 1 1 1 1, 3, 6 1, 5 1
Pseudotriton montanus Baird (Mud Salamander) 2 2
Pseudotriton ruber (Latreille) (Red Salamander) 2 1
Stereochilus marginatus (Hallowell) (Many-lined Salamander)B
Necturus punctatus (Gibbes) (Dwarf Waterdog)A
Pseudobranchus striatus (Le Conte) (Dwarf Siren)B
Siren intermedia Barnes (Lesser Siren) 1 1, 5 1, 5 1 1 1 2 2
Siren lacertina Osterdam (Greater Siren) 2 1, 5
Crocodilians
Alligator mississippiensis (Daudin) (American Alligator) 1 1 1, 5 1 1 2 1 1 1 1 1, 5
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Species AC AP A BH BC FT GR HC/OIS MF PS/S T
Lizards
Anolis carolinensis (Voigt) (Green Anole) 1 1 1, 5 1, 5 1, 5 1 1, 5 2, 3 1, 3, 6 1, 5 1, 3
Ophisaurus attenuatus Cope (Slender Glass Lizard) 5 2
Ophisaurus compressus Cope (Island Glass Lizard)B
Ophisaurus mimicus Palmer (Mimic Glass Lizard)A
Ophisaurus ventralis (Linnaeus) (Eastern Glass Lizard) 4 1 1
Sceloporus undulatus (Bosc and Daudin) (Eastern Fence Lizard) 1 1 1, 5 1, 5 1, 5 1 1 1, 3 1, 3 1 1, 3, 5
Plestiodon inexpectatus (Taylor) (Southeastern Five-lined Skink) 1 1, 5 1 1, 3 1 5 1, 5
Plestiodon laticeps (Schneider) (Broad-headed Skink) 1 1 1, 5 1 1 1 1 1 1, 3, 6 1 1
Plestiodon fasciatus (L.) (Common Five-lined Skink) 5 1 2 1 1 1 1
Plestiodon egregius Baird (Mole Skink) 1 1, 3
Scincella lateralis (Say) (Little Brown Skink) 1 1 1, 5 1 1, 5 1 1 1, 3 1, 3 1 1, 3, 5
Aspidoscelis sexlineata (L.) (Six-lined Racerunner) 1 1 1 1 1 2 1 1, 3 1, 3, 6 1 1, 3
Snakes
Nerodia taxispilota (Holbrook) (Brown Watersnake) 1 1, 6 1 2 2 1 1 1, 6 1 1
Nerodia erythrogaster (Forster) (Plain-bellied Watersnake) 2 1, 5 1 1 2 1 2 1, 6 1 2
Nerodia fasciata (L.) (Southern Watersnake) 1 1 1, 6 1 1, 5 1 1 1 1, 6 1 1
Seminatrix pygaea (Cope) (Black Swampsnake)A
Regina rigida (Say) (Glossy Crayfish Snake) 2 4 5
Rhadinaea flavilata (Cope) (Pine Woods Littersnake) 5
Virginia striatula (L.) (Rough Earthsnake) 5 2 5 1 1, 6 1, 5
Virginia valeriae Baird and Girard (Smooth Earthsnake)B
Storeria dekayi (Holbrook) (Dekay's Brownsnake) 2 2 2 2
Storeria occipitomaculata (Storer) (Red-bellied Snake) 1 1 1, 3, 6 2, 3
Thamnophis sirtalis (L.) (Common Gartersnake) 1 5 1 1 1, 5 1
Thamnophis sauritus (L.) (Eastern Ribbonsnake) 5 1 1 1
Farancia erytrogramma (Palisot de Beauvois) (Rainbow Snake) 1, 6 2 1, 5 1 1
Farancia abacura (Holbrook) (Red-bellied Mudsnake) 1 1 1
Diadophis punctatus Baird and Girard (Ring-necked Snake) 1 1 1 1 1 1, 6 1 5
Cemophora coccinea (Blumenbach) (Scarletsnake) 2 1 2 2 2
Opheodrys aestivus (L.) (Rough Greensnake) 1 4 6 2 1 1 1, 6 1 1
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Species AC AP A BH BC FT GR HC/OIS MF PS/S T
Heterodon platirhinos Latreille (Eastern Hog-nosed Snake) 1 4 1 1 2 4 1 1 1, 3 1, 5 1, 3
Heterodon simus (L.) (Southern Hog-nosed Snake) 2, 5
Coluber constrictor L. (North American Racer) 1 1 1 1 1 1 1 1 1, 3 1 1, 3
Coluber flagellum Shaw (Coachwhip) 1 1 1, 5 1, 5 1, 5 1, 3 1, 3 2 1, 3
Drymarchon couperi (Holbrook) (Eastern Indigo Snake) 2 2 5 1 1, 5 2, 3 1, 3, 6 2, 5 1, 5
Pituophis melanoleucus (Daudin) (Pinesnake) 2 4 1, 6 2, 3 2, 3
Pantherophis alleghaniensis (Holbrook) (Eastern Ratsnake) 1 1 1, 5 1 1, 5 1 1, 5 1 1, 3, 6 1, 5 1, 3
Pantherophis guttatus (L.) (Red Cornsnake) 1 1 1 1 1 1 1 1, 6 1 1
Lampropeltis getula (L.) (Eastern Kingsnake) 1 1, 5 1 1 5 1 5
Lampropeltis elapsoides (Holbrook) (Scarlet Kingsnake) 1 1, 5 1 5 1 1, 6 1
Tantilla coronata Baird and Gerard (Southeastern Crowned 1 1 2, 3 1, 3
Snake)
Micrurus fulvius (L.) (Harlequin Coralsnake) 2 1, 5 1 1, 6 6 2, 6 1
Agkistrodon contortrix (L.) (Copperhead) 1, 5 1 1
Agkistrodon piscivorus (Lacépède) (Cottonmouth) 2, 5 1 1 2 4
Sistrurus miliarius (L.) (Pygmy Rattlesnake) 1 2 6 1, 5 2
Crotalus horridus Linnaeus (Timber Rattlesnake) 1 2, 6 4
Crotalus adamanteus (Palisot de Beauvois) (Eastern Diamond- 2 4 1 1, 5 4 1 1 1 1, 3, 6 1, 5 1, 3
backed Rattlesnake)C
Turtles
Gopherus polyphemus (Daudin) (Gopher Tortoise) 1 1 1 1 1 1 1 1 1 2 1
Kinosternon subrubrum (Lacépède) (Eastern Mud Turtle) 1 1, 5 1 1 1, 6 1, 5
Kinosternon baurii (Garman) (Striped Mud Turtle) 2 2 1, 5 2 1 1 1 1, 6 1 1, 5
Sternotherus odoratus (Latreille) (Eastern Musk Turtle) 2, 5 2 1 2 1 1 1, 6 2
Sternotherus minor (Agassiz) (Loggerhead Musk Turtle) 1 2 2 1, 5 1 2
Trachemys scripta (Schoepff) (Yellow-belled Slider) 1 1 1, 6 1 1 1 1, 5 1 1 1 1
Pseudemys concinna (Le Conte) (River Cooter) 1 2 2 1 2 1 1 1 1, 5
Pseudemys floridana (Le Conte) (Coastal Plain Cooter) 1 1 1 1 1 1 1
Deirochelys reticularia (Latreille) (Chicken Turtle) 1 2 4 1
Terrapene carolina (L.) (Eastern Box Turtle) 1 1 1 1 1 1 1 1, 5 1 1 1
Clemmys guttata (Schneider) (Spotted Turtle) 2 2 1, 6 2 2
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Wildlife Management Area, Natural Area, or Preserve
Species AC AP A BH BC FT GR HC/OIS MF PS/S T
Chelydra serpentina (L.) (Snapping Turtle) 1 1 1 1 1 1 1 1 1, 6 1 1
Apalone ferox (Schneider) (Florida Softshell) 2 1 1, 5 1 1 1 1 1
Apalone spinifera (LeSueur) (Spiny Softshell) 2 1 2 2 1 1 1 1, 5 1
ASpecies on the hypothetical species list for the Altamaha River region that have not been documented from any of our conservation land survey sites or
from any site within 5 km of the Altamaha River main stem.
BSpecies that were not documented from any of our conservation land survey sites but that are known from within 5 km of the Altamaha River main stem
based on museum records.
CAll records from personal communications are from 2008–2016, except the Eastern Diamond-backed Rattlesnake record for Altama Plantation WMA
(2000).