Regular issues
Monographs
Special Issues



Southeastern Naturalist
    SENA Home
    Range and Scope
    Board of Editors
    Staff
    Editorial Workflow
    Publication Charges
    Subscriptions

Other EH Journals
    Northeastern Naturalist
    Caribbean Naturalist
    Neotropical Naturalist
    Urban Naturalist
    Eastern Paleontologist
    Journal of the North Atlantic
    Eastern Biologist

EH Natural History Home

The Herpetofauna of Conservation Lands along the Altamaha River, Georgia
Dirk J. Stevenson and Houston C. Chandler

Southeastern Naturalist, Volume 16, Issue 2 (2017): 261–282

Full-text pdf (Accessible only to subscribers.To subscribe click here.)

 

Site by Bennett Web & Design Co.
Southeastern Naturalist 261 D.J. Stevenson and H.C. Chandler 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 Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 262 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. Southeastern Naturalist 263 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 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. Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 264 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. Southeastern Naturalist 265 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 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 Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 266 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 Southeastern Naturalist 267 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 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. Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 268 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. Southeastern Naturalist 269 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 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 Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 270 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. Southeastern Naturalist 271 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 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 Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 272 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). Southeastern Naturalist 273 D.J. Stevenson and H.C. Chandler 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. Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 274 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. Literature Cited Bauder, J.M., D.J. Stevenson, C.S. Sutherland, and C.L. Jenkins. 2017. Occupancy of potential overwintering habitat on protected lands by two imperiled snake species in the Coastal Plain of the southeastern United States. Journal of Her petology 51:73–88. Beamer, D.A., and T. Lamb. 2008. Dusky salamanders (Desmognathus, Plethodontidae) from the Coastal Plain: Multiple independent lineages and their bearing on the molecular phylogeny of the genus. Molecular Phylogenetics and Evoluti on 47:143–153. Bishop, D.C., and C.A. Haas. 2005. Burning trends and potential negative impacts on flatwoods salamanders. Natural Areas Journal 25:290–294. Buhlmann, K.A., T.S.B. Akre, J.B. Iverson, D. Karapatakis, R.A. Mittermeier, A. Georges, A.G.J. Rhodin, P.P. van Dijk, and J.W. Gibbons. 2009. A global analysis of tortoise and freshwater turtle distributions with identification of priority conservation areas. Chelonian Conservation and Biology 8:116–149. Butler, H., C.D. Camp, G.C. Greer, J.B. Jensen, D. Olive, D.J. Stevenson, and K.M. Stohlgren. 2012. New county records of amphibians and reptiles in Georgia, USA. Herpetological Review 43:453–456. Chandler, H.C., A.L. Rypel, Y. Jiao, C.A. Haas, and T.A. Gorman. 2016. Hindcasting historical breeding conditions for an endangered salamander in ephemeral wetlands of the southeastern USA: Implications of climate change. PLoS ONE 1 1:e0150169. Chandler, H.C., D.J. Stevenson, J.D. Mays, B.S. Stegenga, W.H. Vaigneur, and M.D. Moore. In press. A new trap design for catching small Emydid and Kinosternid turtles. Herpetological Review. Crother, B.L. 2012. Scientific and standard English names of amphibians and reptiles of North America north of Mexico, with comments regarding confidence in our understanding, 7th Edition. Society for the Study of Amphibians and Reptiles Herpetological Circular No. 39. 92 pp. Dallmeyer, D.G., and J. Ray. 2012. Altamaha: A River and Its Keeper. University of Georgia Press, Athens, GA. 187 pp. Diemer, J.E., and D.W. Speake. 1983. The distribution of the Eastern Indigo Snake, Drymarchon corais couperi, in Georgia. Journal of Herpetology 17:256–264. Dorcas, M.E., and W. Gibbons. 2008. Species account: Cottonmouth, Agkistrodon piscivorus. Pp. 427–429, In J. Jensen, C. Camp, W. Gibbons, and M. Elliott (Eds.). Amphibians and Reptiles of Georgia. University of Georgia Press, Athens, GA. 575 pp. Droege, S., A. Cyr., and J. Larivée. 1998. Checklists: An under-used tool for the inventory and monitoring of plants and animals. Conservation Biology 12:1134–1138. Edwards, L., J. Ambrose, and L.K. Kirkman. 2013. The Natural Communities of Georgia. University of Georgia Press, Athens, GA. 675 pp. Southeastern Naturalist 275 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 Enge, K.M., D.J. Stevenson, M.J. Elliott, and J.M. Bauder. 2013. The historical and contemporary distribution of the Eastern Indigo Snake (Drymarchon couperi). Herpetological Conservation and Biology 8:288–307. Erwin, K.J., H.C. Chandler, J.G. Palis, T.A. Gorman, and C.A. Haas. 2016. Herpetofaunal communities in ephemeral wetlands embedded within Longleaf Pine flatwoods of the Gulf Coastal Plain. Southeastern Naturalist 15:431–447. Foster, M.S. 2012. Standard techniques for inventory and monitoring. Pp. 205–272, In R.W. McDiarmid, M.S. Foster, C. Guyer, J.W. Gibbons, and N. Chernoff (Eds.). Reptile Biodiversity: Standard Methods for Inventory and Monitoring. University of California Press, Berkeley, CA. 412 pp. Gibbons, J.W., and R.D. Semlitsch. 1991. Guide to the Reptiles and Amphibians of the Savannah River Site. University of Georgia Press, Athens, GA. 144 pp. Gore, P.J.W., and W. Witherspoon. 2013. Roadside Geology of Georgia. Mountain Press, Missoula, MT. 347 pp. Graham, S.P., E.K. Timpe, and L.R. Laurencio. 2010. Status and possible decline of the Southern Dusky Salamander (Desmognathus auriculatus) in Georgia and Alabama, USA. Herpetological Conservation and Biology 5:360–373. Guyer, C., M.A. Bailey, and R.H. Mount. 2015. Turtles of Alabama. University of Alabama Press, Tuscaloosa, AL. 267 pp. Heyer, W.R., M.A. Donnelly, R.W. McDiarmid, L.C. Hayek, and M.S. Foster. 1994. Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press. Washington DC. 364 pp. Hinkle N.C., C.L. Smith, and C. Bates. 2012. Emergence of 13-year cicadas (Hemiptera:Cicadidae) in Georgia during Spring, 2011. Journal Entomological Science 47:366–368. Hobbs, H.H., Jr. 1981. The Crayfishes of Georgia. Smithsonian Institution Press. Washington, DC. 549 pp. Hyslop, N.L. 2007. Movements, habitat use, and survival of the threatened Eastern Indigo Snake (Drymarchon couperi) in Georgia. Ph.D. Dissertation. University of Georgia, Athens, GA. 132 pp. Hyslop, N.L., J.M. Meyer, R.J. Cooper, and D.J. Stevenson. 2014. Effects of body size and sex of Drymarchon couperi (Eastern Indigo Snake) on habitat use, movements, and home range size in Georgia. Journal of Wildlife Management 78:1–11. Jensen, J., C. Camp, W. Gibbons, and M. Elliott (Eds.). 2008. Amphibians and Reptiles of Georgia. University of Georgia Press, Athens, GA. 575 pp. Jensen, J.B., D.J. Stevenson, S. Graham, M.P. Wallace, Sr., C. Muise, T. Muise, A.B. Safer, F. Snow, R. Preston Jr., G.C. Greer, B. Hudson, and C. Camp. 2011. Additional distributional records of amphibians and reptiles in Georgia, USA. Herpetological Review 42:250–255. Liu, F.R., P.E. Moler, and M.M. Miyamoto. 2006. Phylogeography of the salamander genus Pseudobranchus in the southeastern United States. Molecular Phylogenetics and Evolution 39:149–159. Mazerolle, M.J., L.L. Bailey, W.L. Kendall, J.A. Royle, S.J. Converse, and J.D. Nichols. 2007. Making great leaps forward: Accounting for detectability in herpetological field studies. Journal of Herpetology 41:672–689. Meador, J.R. 2008. The development and evaluation of a freshwater mussel sampling protocol for a large lowland river. M.Sc. Thesis. University of Georgia, Athens, GA. 140 pp. Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 276 Means, D.B. 2000. Southeastern US Coastal Plain habitats of the Plethodontidae: The importance of relief, ravines, and seepage. Pp. 287–301, In R.C. Bruce, R.G. Jaeger, and L.D. Houck (Eds.). The Biology of Plethodontid Salamanders. Kluwer Academic, New York, NY. 485 pp. Means, D.B. 2006. Chapter 6: Vertebrate faunal diversity in Longleaf Pine savannas. Pp. 155–213, In S. Jose, E. Jokela, and D. Miller (Eds.). Longleaf Pine Ecosystems: Ecology, Silviculture, and Restoration. Springer, New York, NY. 438 pp. Means, D.B., and R.C. Means. 2005. Effects of sand pine silviculture on pond-breeding amphibians in the Woodville Karst Plain of north Florida. Pp. 56–61, In W.E. Meshaka Jr. and K.J. Babbitt (Eds.). Amphibians and Reptiles: Status and Conservation in Florida. Krieger Publishing Company, Malabar, FL. 318 pp. Neill, W.T. 1946. Notes on Banded Water Snakes from Georgia. Copeia 1946:255–256. O’Donnell, K.M., and R.D. Semlitsch. 2015. Advancing terrestrial salamander population ecology: The central role of imperfect detection. Journal of Herpetology 49:533–540. Page, L.M., and B.M. Burr. 2011. Peterson Field Guide to Freshwater Fishes of North America North of Mexico. Second Edition. Houghton Mifflin Harcourt, Boston, MA. 663 pp. Palmer, W.M., and A.L. Braswell. 1995. Reptiles of North Carolina. University of North Carolina Press, Chapel Hill, NC. 412 pp. Powell, R., R. Conant, and J.T. Collins. 2016. Peterson Field Guide to Reptiles and Amphibians of Eastern and Central North America, 4th Edition. Houghton Mifflin Harcourt, Boston, MA. 494 pp. Reese, E.G., and D.P. Baxter. 2007. Do invertebrate communities in floodplains change predictably along a river’s length? Freshwater Biology 52:226–239. Roberts, R.L., P.F. Donald, and R.E. Green. 2007. Using simple species lists to monitor trends in animal populations: New methods and a comparison with independent data. Animal Conservation 10:332–339. Schlimm, E.M. 2013. The influence of habitat structure on squamate species richness in the coastal plain of Georgia. M.Sc. Thesis. University of Georgia, Athens, GA. 146 pp. Smith, L.L., W.J. Barichivich, J.S. Staiger, K.G. Smith, and C.K. Dodd Jr. 2006a. Detection probabilities and site occupancy estimates for amphibians at Okefenokee National Wildlife Refuge. American Midland Naturalist 155:149–161. Smith, L.L., D.A. Steen, J.M. Stober, M.C. Freeman, S.W. Golladay, L.M. Conner, and J. Cochrane. 2006b. The vertebrate fauna of Ichauway, Baker County, Georgia. Southeastern Naturalist 5:599– 620. Stevenson, D.J. 1999. The herpetofauna of Fort Stewart, Georgia: Habitat occurrence, status of protected and rare species, and species diversity. Unpublished report. Fort Stewart Fish and Wildlife Branch, Fort Stewart, GA. 71 pp. Stevenson, D.J., and J. Stackhouse. 2012. The amphibians and reptiles of the Altamaha River, Georgia. IRCF Reptiles and Amphibians 19:170–186. Stevenson, D.J., K.J. Dyer, and B.A. Willis-Stevenson. 2003. Survey and monitoring of the Eastern Indigo Snake in Georgia. Southeastern Naturalist 2:393–408. Stevenson, D.J., J.G. Palis, A.D. Walde, A.B. Safer, and C.J. Thawley. 2009a. New distributional records for Georgia reptiles and amphibians. Herpetological Review 40:247–249. Stevenson, D.J., K.M. Enge, L. Carlile, K.J. Dyer, T.M. Norton, N.L. Hyslop, and R.A. Kiltie. 2009b. An Eastern Indigo Snake (Drymarchon couperi) mark–recapture study in southeastern Georgia. Herpetological Conservation and Biology 4:30–42. Stevenson, D.J., G. Beaton, and M.J. Elliott. 2009c. Distribution, status, and ecology of the Say’s Spiketail (Cordulegaster sayi) in Georgia, USA. Bulletin of American Odonatology 11:20–25. Southeastern Naturalist 277 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 Stevenson, D.J., K.M. Stohlgren, E.M. Schlimm, J. Bauder, M.P. Wallace Sr., and W. Taylor. 2011. Distribution records of amphibians and reptiles in Georgia. Herpetological Review 42:255–257. Stevenson, D.J., J. Parker, and M.P. Wallace Sr. 2012. Geographic distribution: Pseudemys concinna concinna (Eastern River Cooter). Herpetological Review 43:442–443. Stevenson, D.J., J.B. Jensen, E.A. Schlimm, and M. Moore. 2015. The distribution, habitat use, activity, and status of the Spotted Turtle (Clemmys guttata) in Georgia. Chelonian Conservation and Biology 14:136–142. Stohlgren, K.M., S.F. Spear, and D.J. Stevenson. 2015. A status assessment and habitat model for the Eastern Diamondback Rattlesnake (Crotalus adamanteus) in Georgia. Unpublished report. Georgia Department of Natural Resources, Atlanta, GA. 22 pp. Trauth, S.E., H.W. Robison, and M.V. Plummer. 2004. The Amphibians and Reptiles of Arkansas. University of Arkansas Press, Fayetteville, AR. 421 pp. Tuberville, T.D., J.D. Wilson, M.E. Dorcas, and J.W. Gibbons. 2005. Herpetofaunal species richness of southeastern national parks. Southeastern Naturalist 4:537–569. Ward, J.P., and D.R. Jackson. 2008. Pseudemys concinna (Le Conte 1830)—River Cooter. Pp. 006.1–006.7, In A.G.J. Rhodin, P.C.H. Pritchard, P.P. van Dijk, P.P. Saumure, K.A. Buhlmann, and J.B. Iverson (Eds.). Conservation Biology of Freshwater Turtles and Tortoises: A Compilation Project of the IUCN/SSC Tortoise and Freshwater Turtle Specialist Group. Chelonian Research Monographs No. 5. 7 pp. Wharton, C.H. 1978. The Natural Environments of Georgia. Georgia Department of Natural Resources, Atlanta, GA. 227 pp. Williamson, G.K., and R.A. Moulis. 1979. Survey of reptiles and amphibians on Fort Stewart and Hunter Army Airfield. Unpublished report. US Army contract DACA 21-77-c- 0155, Fort Stewart, GA. 338 pp. Williamson, G.K., and R.A. Moulis. 1994. Distribution of Amphibians and Reptiles in Georgia: Volume 2 – Locality Data. Savannah Science Museum Special Publication No. 3. Savannah, GA. 712 pp. Wolfe, S.H., J.A. Reidenauer, and D.B. Means. 1988. An ecological characterization of the Florida Panhandle. Biological Report 88. US Fish and Wildlife Service, Minerals Management Service, New Orleans, LA. 277 pp. Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 278 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 Southeastern Naturalist 279 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 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 Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 280 Wildlife Management Area, Natural Area, or Preserve 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 Southeastern Naturalist 281 D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 Wildlife Management Area, Natural Area, or Preserve 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 Southeastern Naturalist D.J. Stevenson and H.C. Chandler 2017 Vol. 16, No. 2 282 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).