A Survey of the Amphibians of Savannah National Wildlife

Refuge, South Carolina and Georgia

Author(s): C. Kenneth Dodd Jr. and William J. Barichivich
Source: Southeastern Naturalist, 16(4):529-545.
Published By: Eagle Hill Institute
https://doi.org/10.1656/058.016.0405
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2017
2017 Southeastern Naturalist
SOUTHEASTERN NATURALIST Vol. 16, No. 4
16(4):529–545
C.K. Dodd, Jr. and W.J. Barichivich

A Survey of the Amphibians of Savannah National Wildlife

Refuge, South Carolina and Georgia
C. Kenneth Dodd Jr. 1, 2,* and William J. Barichivich1

Abstract - From 2004 to 2006, we used a variety of sampling techniques to survey the
amphibians of Savannah National Wildlife Refuge (SNWR), a large protected area strad-
dling the lower portions of the Savannah River on the border between South Carolina and
Georgia. We documented 22 amphibian species—15 frogs and 7 salamanders—with a pos-
sible 23rd species present. Species richness was lower than what might be expected from
amphibian field guides of species inhabiting the adjacent Coastal Plain, likely due to a
lack of specialized habitats, such as temporary ponds and upland pine forest. Amphibians
occupied a variety of habitats and appeared tolerant of the mildly acidic and low-oxygen
conditions of many of the wetlands. Although additional species may be found at SNWR,
this initial survey provides a historic baseline for monitoring amphibian populations as
areas adjacent to the refuge are disturbed, the climate changes, and multi-use management
objectives are implemented within refuge boundaries.

Introduction
Although the Atlantic and Gulf Coastal Plains of southeastern North America
contain a high diversity of amphibians (Dodd 1997, Duellman and Sweet 1999),
information on species richness is unavailable, unpublished, or based on anecdotal
information for many of the protected areas within this region. One of the first steps
to understanding amphibian communities is to establish which species occur within
a region. This knowledge is vitally important because amphibians play significant
roles in ecosystem function, particularly as predators of invertebrates, prey for a
vast array of other species, and in nutrient transfer to and from habitats rich in nu-
trients (e.g., ponds) to areas poor in nutrients, such as sandy uplands (Capps et al.
2014, Earl et al. 2011, Hocking and Babbitt 2014).
In response to the apparent decline of amphibians at the end of the 20th century,
the US Geological Survey established the Amphibian Research and Monitoring
Initiative (ARMI) in 2000 to begin comprehensive studies to assess the status of
amphibian species in the US (Corn et al. 2005). The overall research program was
designed to provide resource managers with information necessary for amphibian
conservation and to identify future research needs (e.g., Grant et al. 2013). Among
a series of tasks, ARMI researchers were directed to establish a network of sites
on US federal lands to monitor amphibian status and trends in the distribution
and abundance of individual species and communities (Muths et al. 2005). Five
1
US Geological Survey, Wetland and Aquatic Research Center, 7920 NW 71st Street,
Gainesville, FL 32653. 2Current address - Department of Wildlife Ecology and Conserva-
tion, University of Florida, Gainesville, FL 32611. *Corresponding author - terrapene600@
gmail.com.
Manuscript Editor: Max Nickerson

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C.K. Dodd, Jr. and W.J. Barichivich

National Wildlife Refuges (NWRs) on the Southeastern Coastal Plain were selected
for potential monitoring, including St. Marks and Lower Suwannee in Florida,
Harris Neck and Okefenokee in Georgia, and Savannah in South Carolina and
Georgia. Other reports present the initial results from St. Marks (Dodd et al. 2007),
Okefenokee (Smith et al. 2006b), Harris Neck (Dodd and Barichivich, 2007), and
Lower Suwannee (Dodd et al. 2017) NWRs. Here, we present the results from our
preliminary survey of the amphibians of Savannah NWR, the first such amphibian
survey on the refuge.

Field-site Description
Savannah National Wildlife Refuge (SNWR) (Fig. 1) comprises 13,387 ha
in Georgia (Chatham County) and South Carolina (Jasper County) immediately
upstream along the Savannah River from the city of Savannah, GA. It is part of
the Savannah Coastal Refuges Complex under the jurisdiction of the US Fish and
Wildlife Service’s (USFWS) National Wildlife Refuge system. The refuge has an

Figure1. Map showing

the extent of Savan-
nah National Wildlife
Refuge and the loca-
tion of sampling sites
from 2004 to 2006.
BI = Bear Island,
DT = Dodge Tram
Road, ND = North
(Wildlife) Drive, ST
= Solomon Tract, and
WD = Wildlife Drive.

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C.K. Dodd, Jr. and W.J. Barichivich

extensive history of human occupation and use, from Native Americans through
the plantation era, when the bottomlands and freshwater tidal marshes were
extensively diked and modified for rice production. Dike construction began
in the mid-to-late 1700s, with peak rice harvests occurring from 1850 to 1860
(Rowland 1987, USFWS 2011). Established in April 1927, the refuge is primar-
ily managed for waterfowl, and water levels within the former rice fields (1364
ha) are carefully regulated via 22 water-control structures. The refuge staff occa-
sionally clears vegetation from impounded areas, resulting in a variety of marsh
habitats of different depths, vegetation structure, and species composition. Spe-
cific management objectives seek to provide a refuge and breeding ground for
native birds and other wildlife, protection for endangered and threatened species,
a sanctuary for migratory birds, native habitat maintenance, and educational/rec-
reational activities (USFWS 2011).
The northern part of the refuge (upstream from the impounded freshwater tidal
marshes) consists mostly of extensive tracts of bottomland hardwoods (2975 ha)
and cypress–gum swamp (4726 ha) that may or may not be periodically flooded
(Garman and Nielsen 1992). The habitat classification here and below is as depicted
on Fig. 16 in USFWS (2011). In addition, there are forested islands within the main
Savannah River channel. These islands, the largest of which is Bear Island (312 ha),
contain small channels and an extensive number of woodland pools and “creeks”
that hold water for varying amounts of time. There is only 1 large pond on the ref-
uge (Kingfisher Pond, an old borrow pit; site DT-2 on Fig. 1) not associated with the
bottomland forest. River bluffs and upland terraces on the refuge are few because
the refuge boundary often terminates at the base of the river bluff. However, some
upland hardwoods and forested slopes are present along Dodge Tram Road (sites
DT in Fig. 1) on the north side of the river, and more extensive uplands, including
bottomland forest and a very small area of oak hammock, are found on the south
side of the river east of O’Leary (as marked on the USGS 7.5’ Port Wentworth topo-
graphical map). This 403-ha parcel, the Solomon Tract (sites ST on Fig. 1), was one
of the most recent additions to SNWR when we began our study. We sampled this
area in connection with the USFWS malformed frog survey (Reeves et al. 2013).
Additional information on the climate and vegetative communities of SNWR is
provided in USFWS (2011).

Methods
We used a variety of methods to sample amphibians at SNWR (Appendix 1)
from 2004 to 2006. Terrestrial sampling techniques focused on diurnal time-con-
strained field searches (i.e., visual encounter surveys [VES]) in the open and under
cover-objects (e.g., logs, rocks, leaf litter). We conducted terrestrial surveys along
Dodge Tram Road in bottomland wetland forest and in upland hardwoods on the
slopes, as well as in the Solomon Tract. Most terrestrial VES lasted 20–30 min per
site, and were conducted by 2–4 observers. Aquatic sampling employed dip nets
(Memphis Net and Twine Co., HDD-2 with 4.7 mm [3/16”] sq. Delta mesh), cray-
fish traps (Johnson and Barichivich 2004), aural surveys (incidental observations
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C.K. Dodd, Jr. and W.J. Barichivich

while conducting VES and nocturnal road cruising), and visual-encounter obser-
vations. Typically, we deployed 2–4 crayfish traps per site per night depending
on water availability. Aquatic sampling focused on the impounded wetlands and
adjacent freshwater marshes, at Kingfisher Pond, and in the few small, shallow
temporary pools located in the Solomon Tract.
We sampled 28 sites in 2004 (25 wetlands, 3 terrestrial) in April, June/July,
and October; we deployed crayfish traps for 60 trap nights. Terrestrial sampling
focused on upland areas on Bear Island (along a 470-m survey transect previously
established by the USFWS), the Solomon Tract, and along Dodge Tram Road. In
2005, we sampled 10 sites (4 wetland, 6 terrestrial) in March, 15 sites in May (11
wetlands, 4 terrestrial, including several additional sites on Bear Island), and 9 sites
in July (5 wetland, 4 terrestrial). We only sampled 10 wetland sites in 2006.
In addition to sampling, we examined Jasper and Chatham county museum re-
cords from the American Museum of Natural History (New York, NY), Carnegie
Museum (Pittsburgh, PA), Charleston Museum (Charleston, SC), Florida Museum
of Natural History (Gainesville, FL), Francis Marion University (Jeffrey Camper;
Florence, SC), Georgia Museum of Natural History (Athens, GA), Georgia South-
ern University (Statesboro, GA), HerpMapper (https://www.herpmapper.org/),
iNaturalist (https://www.inaturalist.org/), Museum of Zoology (University of
Michigan; Ann Arbor, MI), North Carolina State Museum (Raleigh, NC), Savannah
Science Museum (Williamson and Moulis 1994), and US National Museum (Wash-
ington, DC). No amphibians originating from SNWR were found in any of these
holdings. The South Carolina Herpetological Atlas contains 2 amphibian records
from SNWR (Hyla cinerea [Green Treefrog], Lithobates grylio [Pig Frog]) with
several additional species found in close proximity (Ambystoma opacum [Marbled
Salamander], Eurycea cirrigera Green [Southern Two-lined Salamander], Anaxyrus
terrestris [Southern Toad], Hyla chrysoscelis [Cope’s Gray Treefrog], H. femoralis
[Pine Woods Treefrog], H. squirella [Squirrel Treefrog], Lithobates catesbeianus
[American Bullfrog]) southwest of Hardeeville along Purrysburg Road.
We collected standard abiotic field parameters likely to influence amphibian
species distribution at most of the wetlands sampled. Using a Hydrolab® Quanta®
water-quality meter, we measured water temperature (ºC), conductivity (µS/cm),
pH, dissolved oxygen (mg/l), and % dissolved oxygen.

Results
From 2004 to 2006, we detected 22 species of amphibians (15 frogs, 7 sala-
manders) at SNWR, with a possible 23rd species (Siren intermedia [Lesser Siren])
unconfirmed (Appendix 1). The suspected Lesser Siren consisted of 3 individu-
als regurgitated by a Nerodia fasciata L. (Banded Watersnake) that we trapped
in a crayfish trap on 26 May 2006. The state of digestion made identification
impossible because identification as subadult S. lacertina (Greater Siren) could
not be ruled out. We heard 2 additional species, Pseudacris ocularis Bosc and
Daudin [Little Grass Frog]) and P. nigrita Le Conte (Southern Chorus Frog),
calling on 24 March 2005 immediately adjacent to SNWR along Chisolm Road
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(32°13.398'N, 81°5.647'W), which connects Dodge Tram Road with Purrysburg

Road north–northeast of the refuge. The most amphibian species-rich wetland
was Kingfisher Pond (n = 12; site DT-2), where we also observed huge numbers
of Lithobates heckscheri (River Frog) tadpoles in late July 2005 (662 tadpoles in
4 traps on 28 July; 470 the next day). Tadpoles of this species form large schools,
and we observed very large numbers of tadpoles gulping air even as we set and
picked up crayfish traps. Although we found few species in the rice fields, these
fields provide extensive habitat (1364 ha) for ranid frogs and aquatic salaman-
ders (Amphiuma, Siren); these are the only areas where we found Greater Sirens
and Amphiuma means (Two-toed Amphiuma). We detected no amphibians at site
ND-5, an area along the Little Black River subject to extreme tidal fluctuation
(2.7–4.5 m). We set traps at site WD-8 for 7 d in May and July 2005, but observed
only a single Lithobates sp. at this open-marsh site; however, we trapped Two-
toed Amphiumas and Pig Frogs at nearby site WD-5.
We collected water-quality field parameters at 13 locations from 2004 to 2006,
with multiple ponds sampled more than once or twice (Table 1). Water temperatures
were generally warm (15.3–26.4 ºC) because we made all observations between
March and October. The lowest conductivities, ≤100 μS/cm, were measured at
sites located outside the river floodplain (DT-2, ST-2, ST-3, ST-5). Although these
wetlands range in size and origin from tiny woodland pools to a 6-ha borrow pit,
they appear to share a rainwater-dominated hydrologic regime. The remaining sites
are in the flooplain and exhibited greater conductivities than those in uplands. With
the exception of WD-5, these sites are within an agricultural area—historically rice
fields—defined by a series levees and canals. While geographically nearby, WD-5
is located in a natural floodplain. The freshwater tidal influence of the Savannah
River and its many tributaries may account for the general increase in conductivity;
however, locations WD-3 and WD-8 were >1000 μS/cm. Sample locations tended
to be neutral (pH 7.01) to slightly acidic (pH 6.10), with very little variation be-
tween sites and sampling events. Dissolved oxygen levels, both total and %, were
low and variable across most sites. Although inadequate for most fish, 3 woodland
pools, ST-2, ST-3, and ST-5, were among the most oxygen rich (>2 mgL-1 or 20%)
of all the sites sampled.

Discussion
Publications suggest that 43 amphibians (17 salamanders and 26 frogs) might
be found within the general vicinity of SNWR (Dorcas and Gibbons 2008, Jensen
et al. 2008, Mitchell and Gibbons 2010). Because of specific habitat requirements
and a dearth of uplands on the refuge, we would not expect many of these species
to occur on site (e.g., temporary pond breeders such as, Ambystoma cingulatum
Cope [Frosted Flatwoods Salamander], A. mabeei Bishop [Mabee’s Salamander],
A. tigrinum Green [Eastern Tiger Salamander], Hyla gratiosa LeConte [Barking
Treefrog], Pseudacris ornata Holbrook [Ornate Chorus Frog], Lithobates capito
LeConte [Gopher Frog]), nor should some species, if present, be expected to occur
widely within its boundaries (e.g., Hyla avivoca Viosca [Bird-voiced Treefrog]).
533
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Table 1. Water-quality data collected during amphibian sampling at Savannah National Wildlife Refuge from 2004 to 2006, supplemented by data collected
by the USGS National Water Quality Laboratory from 2006 to 2008. Means followed by standard deviations in parentheses. SpC = specific conductivity,
% DO = percent dissolved oxygen.

Water SpC DO
Site 07/01/04 10/21/04 03/24/05 05/20/05 07/28/05 05/25/06 08/16/06 temp (°C) (μS/cm) (mg/L) pH % DO
DT-2 X X X X X X X 23.4 (3.8) 37 (12) 1.38 (1.57) 6.10 (0.78) 14.9 (15.9)
HQ-1 X X X X X 25.2 (2.5) 513 (212) 0.81 (0.48) 6.32 (0.36) 10.0 (6.2)
ND-3 X X X X 26.4 (1.8) 279 (112) 0.88 (0.46) 6.48 (0.36) 10.7 (5.5)

534
ND-4 X X X X 24.5 (1.2) 477 (170) 0.89 (0.62) 6.25 (0.36) 10.6 (7.2)
ST-2 X X 23.5 (3.9) 66 (13) 3.21 (0.13) 6.90 (0.01) 35.5 (3.5)
ST-3 X X 21.2 (1.4) 100 (8) 2.80 (1.13) 6.57 (0.55) 31.7 (13.4)
ST-5 X 20.8 88 2.02 6.22 22.8
Southeastern Naturalist

WD-3 X X X X X X 23.3 (3.7) 1009 (141) 1.37 (1.11) 6.63 (0.38) 14.6 (11.0)
WD-4 X 20.0 554 0.97 6.70 10.7
C.K. Dodd, Jr. and W.J. Barichivich

WD-5 X 19.3 135 0.66 6.00 7.1

WD-6 X 15.3 853 2.27 7.01 22.8
WD-7 X X 22.6 (3.7) 855 (150) 1.51 (1.07) 6.64 (0.39) 16.5 (11.1)
WD-8 X X 25.4 (2.4) 1915 (85) 0.33 (0.01) 6.28 (0.06) 4.1 (0.3)
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C.K. Dodd, Jr. and W.J. Barichivich

We observed the Little Grass Frog and Southern Chorus Frog within sight of the
refuge boundary in temporary upland pools. Several species (e.g., Southern Two-
lined Salamander, Pseudotriton montanus Baird [Mud Salamander], Stereochilus
marginatus Hallowell [Many-lined Salamander]) may yet be found on the refuge.
Still, the refuge contains substantial amphibian diversity, particularly those species
that occupy the extensive marshlands bordering the lower Savannah River. The bot-
tomlands, cypress–gum swamps, ravine slopes, and islands within the floodplain
offer a diverse suite of habitats where species from the Piedmont uplands may enter
into the lower Coastal Plain.
Most of the amphibians we detected during our surveys are common in the
lower Coastal Plain of South Carolina and Georgia (Dorcas and Gibbons 2008,
Jensen et al. 2008, Mitchell and Gibbons 2010). Of particular interest, however,
is the presence of Ambystoma opacum (Marbled Salamander) on Bear Island and
possibly other islands within the Savannah River. This small (10 cm total length)
terrestrial species deposits eggs under debris during the autumn in shallow depres-
sions (Mitchell and Gibbons 2010). During winter rains, the shallow depressions
fill and form pools in which the larvae hatch and complete development. This spe-
cies is not known to cross open water, particularly large fast-flowing rivers, and
may have colonized the river islands during floods. Alternatively, the species could
have been present in backwater forests (Zone IV of Larson et al. 1981) and became
subsequently isolated as the river changed course through the broad floodplain.
Another unusual inhabitant of this region, both on the river islands and at
Kingfisher Pond, is Anaxyrus fowleri (Hinckley) (Fowler’s Toad). This species is
normally associated with the upland Piedmont Region, and is rarely found in the
Coastal Plain along rivers downstream (Dodd 2013, Floyd 2008). The toad’s entry
into the Coastal Plain was either facilitated by the riparian corridor, or toads could
have been washed downstream from the Piedmont during floods. We did not detect
Acris crepitans Baird (Eastern Cricket Frog), another species thought to enter the
Coastal Plain through riparian corridors (Dodd 2013).
The Bird-voiced Treefrog, enters SNWR only at the northern boundary of the
refuge, where it was heard at the abandoned Chisholm Landing. Other species
(Desmognathus cf. auriculatus [Southern Dusky Salamander], Notophthalmus
viridescens [Eastern Newt], Plethodon variolatus [South Carolina Slimy Sala-
mander], Gastrophryne carolinensis Holbrook [Eastern Narrow-mouthed Frog],
Pine Woods Treefrog, River Frog) are not widely distributed throughout the
refuge because of limited available habitat. These species appear to be common
where found, however, and all are widely distributed outside the refuge bound-
aries, with the exception of Southern Dusky Salamander (Graham et al. 2010,
Maerz et al. 2015).
Mention should be made concerning the Desmognathus (dusky salamanders)
found within SNWR. We observed dusky salamanders in the bottomland forest
among the seeps and swampy areas west of Dodge Tram Road in South Carolina
(Zones IV and V of Larson et al. 1981) and in the upland forest of the Solomon
Tract on the Georgia side of the Savannah River. The systematics of Desmognathus

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in the Coastal Plain, especially along the major rivers, is by no means clear. Coastal
Desmognathus may be partitioned into a variety of clades that do not appear to fol-
low clear biogeographic patterns (Beamer and Lamb 2008). In the lower portions of
the Plain in South Carolina and Georgia, Desmognathus can be partitioned into nu-
merous lineages, some more closely related to distant species (D. conanti Rossman
[Spotted Dusky Salamander], D. fuscus Rafinesque [Northern Dusky Salamander],
D. monticola Dunn [Seal Salamander]) than to nearby conspecifics. It may be that
there are numerous morphologically cryptic species in this region (Beamer and
Lamb 2008, Means et al. 2017), perhaps partitioned by habitat preferences. For
the interim, we classify SNWR dusky salamanders as D. cf. auriculatus [Southern
Dusky Salamander] until relationships are resolved.
As noted by Yu et al. (2015), water-quality information reflects watershed envi-
ronments and is valuable for assessing the suitability of wetlands for amphibians.
Our preliminary water-quality indicators were typical of what might be expected
for this region. Wetlands were generally slightly acid to neutral in pH, reflecting
the tannic acid nature of the bottomland forest and the highly organic impounded
wetlands. Many amphibians can tolerate rather acidic environments (references in
Dodd 2013), and the pH readings we observed were well within the tolerance limits
of these species-based laboratory observations (Freda 1986). With few exceptions,
oxygen saturation tended to be low, again reflecting the highly eutrophic nature of
the impounded wetlands exposed to intensive solar radiation, or to the small size
of the nearly stagnant woodland pools. However, these values are also well within
the range for field studies of anurans elsewhere (e.g., Dodd and Barichivich 2007,
Noland and Ultsch 1981, Wassersug and Seibert 1976). Frogs frequently are found
in waters with low DO, leading tadpoles to gulp air, especially during the latter
stages of development when water temperatures are high in summer (Wassersug
and Seibert 1976). This behavior likely facilitates long larval periods, particularly
of ranids, in SNWR’s open canopied impounded wetlands and marshes.
SNWR has not been the object of intensive land use after the bottomland forest
was cleared for rice plantations in the 1700s and 1800s. Taxodium distichum (L.)
Rich (BaldCypress) and other forest species were harvested throughout the ripar-
ian corridor, but today, this region is covered by dense 2nd- and 3rd-growth forest
that is managed for hunting and other recreational use. Timber harvest is currently
absent from the refuge, although pine plantations occur to the east of Dodge Tram
Road. Urban development occurs along the western edge of the Solomon Tract right
up to the border of the refuge. The lower impounded marshlands are managed for
waterfowl, with occasional dredging that may adversely affect amphibians (Aresco
and Gunzburger 2004). Access to many areas of the refuge is strictly regulated, with
most human use occurring along a dirt road that allows visitors to view wildlife
through the marshes, fishing along the river and at Kingfisher Pond, and seasonal
hunting in some areas. These activities appear consistent with the maintenance of
the amphibian community, despite occasional disruption from dredging.
In general, protected areas contain only a fraction of amphibian biodiver-
sity (Nori et al. 2015). However, it appears that many protected areas on the

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C.K. Dodd, Jr. and W.J. Barichivich

Southeastern Coastal Plain contain a substantial number of amphibians (Table 2),

except for some of the rarer species such as the 2 species of flatwoods salaman-
ders and Southern Dusky Salamander. Plans to enlarge the refuge, as detailed in
USFWS (2011), would add upland habitats that might contain additional species
and expand the limited distribution of upland amphibians currently present. Ex-
pansion would also allow for the protection of core habitat for wetland-breeding
amphibians (Semlitsch 2000, Semlitsch and Jensen 2001). Monitoring the SNWR
amphibian community would enable biologists to assess the potential impacts
of future environmental changes (e.g., sea-level rise, pollution, climate change,
urban and silviculture encroachment; USFWS 2011) and management practices
(e.g., prescribed fire, herbicide spraying, fluctuating water levels) (Lindenmayer
and Likens 2010).

Acknowledgments
We thank the staff of SNWR for supporting and permitting our work at the refuge.
ARMI research and monitoring activities at SNWR were conducted under USFWS Special
Use Permits No. 41620-04016 and 05-41650-4. We thank Steve Bennett, Linda Casey, Ju-
lia Earl, Denise Gregoire, Joe Mitchell, and Clint Smith for assisting in some of the field
surveys. We are grateful to Kevin Smith and Anna Tutterow for supplying South Carolina

Table 2. Amphibian species richness on selected protected private, federal, and state lands on the
Atlantic Coastal Plain of the Southeastern US (exclusive of peninsular Florida). NF = national forest,
NP = national park, NS = national seashore, NWR = national wildlife refuge, SF = state forest, and
WMA = wildlife management area.

Location Area(ha) Salamanders Frogs Source

Alligator River NWR 69,091 2 15 Meyers and Pike (2006)
Altamaha River corridorA 58,208 18 23 Stevenson and Chandler (2017)
Big Bend WMA 29,111 10 19 Enge and Wood (1998), Surdick (2013)
Blackwater SFB - 6 10 Enge (2002)
Congaree NP 10,633 6 21 Tuberville et al. (2005)
Cumberland Island NS 36,415 5 12 Shoop and Ruckdeschel (2000)
Francis Beidler Forest 1398 8 16 Hall (1994)
Francis Marion NF 101,000 8 17 Schurbon and Fauth (2003)
Great Dismal SwampC 57,469 8 19 Delzell (1979)
Harris Neck NWR 1285 1 12 Dodd and Barichivich (2007)
Ichauway Plantation 11,300 13 8 Smith et al. (2006a)
Jug Bay Wetlands Sanctuary 200 7 12 Smithberger and Swarth (1993)
Okefenokee NWR 182,727 16 21 Smith et al. (2006b)
Roanoke River corridorD 8500 7 12 Lamb et al. (1998)
Savannah NWR 13,387 7 15 This paper
Savannah River Site 198,344 17 26 Gibbons and Semlitsch (1991)
St. Marks NWR 27,500 13 21 Dodd et al. (2007), ongoing USGS
surveys
A
Surveys of 2 private preserves and 11 Georgia Wildlife Management Areas.
B
Surveys of 2 small seepage bogs; area not given.
C
Great Dismal Swamp NWR and Dismal Swamp State Park.
D
A series of 5 preserves jointly managed by The Nature Conservancy and Georgia-Pacific.

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Heritage Program database records from Jasper County. Russ Webb and John Robinette of
SNWR facilitated work on the refuge, and we greatly appreciate their assistance. Research
was carried out under US Geological Survey animal care and use protocols for sampling
and handling amphibians. This study was funded by the USGS Amphibian Research and
Monitoring Initiative (ARMI). Any use of trade, product, or firm names is for descriptive
purposes only and does not imply endorsement by the US Government. This is contribution
number 603 of the US Geological Survey’s Amphibian Research and Monitoring Initiative.
Additional water-quality data are available from the National Water Information System
(water data.usgs.gov/nwis) and the biological and water-quality data used herein can be
found at doi.org/10.5066/F7ST7NRD.

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 Appendix 1. Summary table of amphibians observed or collected during surveys at Savannah National Wildlife Refuge from 2004 to 2006. Habitats: (F) 2017
bottomland hardwood forest; (S) ravine slope between bottomland hardwood and upland forests consisting of mixed slope forest and low, swampy, spring-
fed seeps; (P) large manmade permanent pond (Kingfisher Pond); (T) small woodland pool; (W) extensive freshwater marshes that were former rice fields;
(U) upland deciduous forest. Sampling: (V) visual-encounter surveys, including litter searches, incidental captures, and detection of frog calls; (A) aquatic
sampling using crayfish traps and dipnets. -An unidentified Lithobates was observed at WD-8.

Site
Bear
Isl. BI-1 BI-4 BI-5 BI-6 BI-7 DT-1 DT-2 DT-3 DT-4 DT-5 DT-6 HQ-1 ND-1 ND-2 ND-3 ND-4 ND-5
Habitat F F F F F F S P S S S S W W W W W W
Sampling V V V V V V V V V V V V A A A A A A
Species
Acris gryllus (Le Conte) X X X X X X
(Southern Cricket Frog)
Anaxyrus fowleri (Hinckley) X X

542
(Fowler’s Toad)
A. terrestris (Bonnaterre) X X X X X
(SouthernToad)
Gastrophryne carolinensis (Holbrook) X X X X
Southeastern Naturalist

(Eastern Narrow-mouthed Frog)

Hyla avivoca Viosca X X
C.K. Dodd, Jr. and W.J. Barichivich

(Bird-voiced Treefrog)
Hyla chrysoscelis Cope X X X X X X X X X
(Cope’s Gray Treefrog)
H. cinerea (Schneider) X X X X
(Green Treefrog)
H. femoralis Bosc X X X X
(Pine Woods Treefrog)
H. squirella Bosc X X X
(Squirrel Treefrog)
Lithobates catesbeianus (Shaw) X X X
Vol. 16, No. 4

(American Bullfrog)
 Site
2017

Bear
Isl. BI-1 BI-4 BI-5 BI-6 BI-7 DT-1 DT-2 DT-3 DT-4 DT-5 DT-6 HQ-1 ND-1 ND-2 ND-3 ND-4 ND-5
L. clamitans (Latreille) X X X X X X X X X
(Green Frog)
L. grylio (Stejneger) X X X X
(Pig Frog)
L. heckscheri (Wright) X
(River Frog)
L. sphenocephalus (Cope) X X X X X X X X X X X
(Southern Leopard Frog)
Pseudacris crucifer (Wied-Neuwied) X X X X
(Spring Peeper)
Amphiuma means Garden X X X
(Two-toed Amphiuma)
Ambystoma opacum (Gravenhorst) X X

543
(Marbled Salamander)
Desmognathus cf. auriculatus (Holbrook) X X X
(Southern Dusky Salamander)
Southeastern Naturalist

Eurycea quadridigitata (Holbrook) X

(Dwarf Salamander)
C.K. Dodd, Jr. and W.J. Barichivich

Notophthalmus viridescens (Rafinesque) X

(Eastern Newt)
Plethodon variolatus (Gilliams)
(South Carolina Slimy Salamander)
Siren intermedia Barnes ?
(Lesser Siren)
S. lacertina Osterdam X X X X
(Greater Siren)
Species count 5 3 3 2 3 1 7 12 8 8 9 5 5? 2 4 5 4 0
Vol. 16, No. 4
 Site 2017

SCL1 ST-1 ST-2 ST-3 ST-4 ST-5 ST-6 WD-1 WD-2 WD-3 WD-4 WD-5 WD-6 WD-7 WD-8 # of sites
Habitat W U T T T U U W W W W W W W W
Sampling A A A A A A A A A A A A A A A
Species
Acris gryllus (Le Conte) X X X 10
(Southern Cricket Frog)
Anaxyrus fowleri (Hinckley) 2
(Fowler’s Toad)
A. terrestris (Bonnaterre) X X 7
(SouthernToad)
Gastrophryne carolinensis (Holbrook) X 5
(Eastern Narrow-mouthed Frog)
Hyla avivoca Viosca 2
(Bird-voiced Treefrog)

544
Hyla chrysoscelis Cope X X X X X 14
(Cope’s Gray Treefrog)
H. cinerea (Schneider) X X X X 8
Southeastern Naturalist

(Green Treefrog)
H. femoralis Bosc 4
C.K. Dodd, Jr. and W.J. Barichivich

(Pine Woods Treefrog)

H. squirella Bosc X X 5
(Squirrel Treefrog)
Lithobates catesbeianus (Shaw) X X 5
(American Bullfrog)
L. clamitans (Latreille) X X X X 13
(Green Frog)
L. grylio (Stejneger) X X X X X X 10
(Pig Frog)
L. heckscheri (Wright) X 2
Vol. 16, No. 4

(River Frog)
 Site
2017

SCL1 ST-1 ST-2 ST-3 ST-4 ST-5 ST-6 WD-1 WD-2 WD-3 WD-4 WD-5 WD-6 WD-7 WD-8 # of sites
L. sphenocephalus (Cope) X X X X X X X X X 20
(Southern Leopard Frog)
Pseudacris crucifer (Wied-Neuwied) X 5
(Spring Peeper)
Amphiuma means Garden X 4
(Two-toed Amphiuma)
Ambystoma opacum (Gravenhorst) X 3
(Marbled Salamander)
Desmognathus cf. auriculatus (Holbrook) X 4
(Southern Dusky Salamander)
Eurycea quadridigitata (Holbrook) X X 3
(Dwarf Salamander)
Notophthalmus viridescens (Rafinesque) X X X X X 6

545
(Eastern Newt)
Plethodon variolatus (Gilliams) X X X X 4
(South Carolina Slimy Salamander)
Southeastern Naturalist

Siren intermedia Barnes 1?

(Lesser Siren)
C.K. Dodd, Jr. and W.J. Barichivich

S. lacertina Osterdam X X 6
(Greater Siren)
Species count 1 4 3 8 8 2 9 3 2 8 2 2 1 3 0*
Vol. 16, No. 4