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The Herpetofauna of Lake Woodruff National Wildlife Refuge, Florida
Terence M. Farrell, Melissa A. Pilgrim, Peter G. May, and W. Boyd Blihovde

Southeastern Naturalist, Volume 10, Issue 4 (2011): 647–658

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2011 SOUTHEASTERN NATURALIST 10(4):647–658 The Herpetofauna of Lake Woodruff National Wildlife Refuge, Florida Terence M. Farrell1,*, Melissa A. Pilgrim2, Peter G. May1, and W. Boyd Blihovde3 Abstract - We monitored the herpetofauna of a large conservation area in peninsular Florida from 1990 to 2010, visiting the refuge on over 2000 different days in that period and using a variety of sampling techniques. Our goals were to evaluate the species richness and abundance of herpetofauna (amphibians and reptiles) on the site. During the course of our study, we documented 72 species (22 amphibian and 50 reptile species); thus, the site has impressive species richness relative to other conservation areas in the southeastern United States. Five of the 72 species (6.9%) we found were introduced species. Continued monitoring of the diverse herpetofauna of Lake Woodruff National Wildlife Refuge could facilitate a better understanding of ongoing changes in species abundances and community assemblages occurring in the ecosystems of peninsular Florida. Introduction Faunal surveys help identify critical locations for conservation efforts and can also provide valuable baseline data for assessing changes in abundance and species composition. Inventory and monitoring studies of herpetofauna (i.e., reptiles and amphibians) can be particularly informative. Herptetofauna often play major roles in ecosystem function as a result of their abundance (e.g., Regester et al. 2006) and importance as both predators and prey (e.g., Davic and Welsh 2004, Dial and Roughgarden 1995). Thus, both reptiles (e.g., Beaupre and Douglas 2009) and amphibians (e.g., Welsh and Droege 2001) are considered important bioindicators of ecosystem health and integrity. The declines detected in both amphibian (Houlahan et al. 2000) and reptile (Gibbons et al. 2000) populations are another reason to closely monitor herpetofauna. Extensive surveys of herpetofauna are only occasionally available for many conservation areas. Recently, however, the Southeast Coast Network of the National Park Service, which includes many refuges along the Atlantic seaboard of the Carolinas, Georgia, and North Florida, inventoried herpetofauna on 16 refuges (Tuberville et al. 2005). Unfortunately, most conservation areas in peninsular Florida have received less study despite their rich herpetofauna and susceptibility to anthropogenic impacts (e.g., introduced species and proximity to human development). In this paper, we present the results of extensive field surveys completed in the last two decades at a major conservation area in peninsular Florida: Lake Woodruff National Wildlife Refuge. We document the 1Biology Department, Stetson University, 421 North Woodland Boulevard, DeLand, FL 32720. 2Division of Natural Sciences and Engineering, University of South Carolina Upstate, 800 University Way, Spartanburg, SC 29303. 3Lake Woodruff National Wildlife Refuge, 2045 Mud Lake Road, DeLeon Springs, fl32130. *Corresponding author - 648 Southeastern Naturalist Vol. 10, No. 4 herpetofauna of this refuge, with the goal of providing information that will inform conservation efforts. Field Site Description Lake Woodruff National Wildlife Refuge (LWNWR) is an 8700-hectare conservation area along the eastern floodplain of the St. Johns River in Volusia County, FL. While LWNWR has little topographic relief (approximately 10 m), a variety of habitats are present. Areas of xeric upland forest, mesic forest, cypress swamp, freshwater marsh, temporary pools, permanent ponds and lakes, and several large, shallow anthropogenic impoundments are found at LWNWR (USFWS 2008). Prior studies of the herpetofauna at this site focused on individual species inhabiting the refuge, such as Terrapene carolina (Common Box Turtle; Pilgrim et al. 1997) or Sistrurus miliarius (Pigmy Rattlesnake; May et al. 1996). Since much of LWNWR is difficult for humans to access, we focused our sampling efforts on the most accessible portions of the refuge that extend from the Mud Lake Road entrance and along the firebreaks connected to the Highland Park Road entrance (Fig. 1). The areas we extensively sampled encompassed roughly 8.4% of the entire refuge (730 of 8700 hectares) and included all major habitat types found in the refuge. Figure 1. A map of the most intensely studied portion of LWNWR. The length of levee sampled in the crepuscular levee surveys runs from point “A” to point “B”. Yellow triangles indicate the location of the four upland drift fence arrays and the red triangles indicate the location of the 13 lowland drift fence arrays. The blue circle indicates Mud Lake Road entrance to the refuge. 2011 T.M. Farrell, M.A. Pilgrim, P.G. May, and W.B. Blihovde 649 Methods Species richness We used a wide variety of techniques (e.g., drift fences, coverboard arrays, auditory surveys, aquatic funnel traps, dip netting, diurnal visual surveys, and nocturnal visual surveys) to determine what reptile and amphibian species occur on LWNWR. We investigated the refuge herpetofauna at all times of year on over 2000 days during the past two decades. From our quantitative data sets and opportunistic sampling efforts we compiled a list of the species observed at LWNWR and indicated the habitat types in which each species was typically found. We also used documented information (unambiguous photographs) gathered from the refuge staff and local naturalists to document the occurrence of rare or cryptic species. Drift fence arrays We installed 17 drift fence arrays during the fall of 2001. We used a linear 15-m x 1-m (length by height) stretch of pre-staked silt fencing in each array. We buried the bottom of the fence 15 cm to prevent organisms from escaping under the fence. We placed 6 funnel traps and 4 pitfall traps along each array. We built funnel traps out of hardware cloth, using a design similar to Fitch (1987). We placed dead vegetation over each funnel trap to minimize heat stress of captured animals and used 18.9-L buckets as pitfall traps. We drilled small holes in the bottom of each bucket to allow for drainage in wet weather. We placed a large sponge in each bucket to reduce desiccation of captured animals in dry weather and prevent drowning of captured animals in wet weather. We buried buckets so that their lips were flush with ground level and a portion of each lip was flush with the fence line. For logistical reasons, we implemented a sampling protocol where each season we opened half of the drift fence arrays for approximately 3 weeks, closed them, and then opened the remaining group of arrays for approximately 3 weeks. We randomly determined the set of arrays that would be checked first during a sampling run. We collected our spring sample from 3/10/02–4/25/02, summer sample from 6/8/02–8/7/02, fall sample from 9/23/02–11/13/02, and winter sample from 1/12/02–3/4/03. We checked open drift fences daily to avoid killing trapped animals. Of the 17 arrays we installed, 4 were located in upland habitats and 13 were located in lowland habitats. The upland sites were dominated by scrub habitat and pine/palmetto flatwoods, while lowland sites were dominated by Spartina bakeri Merr. (Sand Cordgrass), marsh edges, mesic hammock habitat, and cypress swamp. Our seasonal sampling protocol resulted in a total of 1614 array days: 394 array days from upland sites and 1220 array days from lowland sites. We calculated the relative abundance of each species captured within upland and lowland sites as the number of individuals captured per array day. In addition, we calculated the percentage of total snake, lizard, or anuran captures each species represented within the upland and lowland samples. In one sampling period, a large number of Scaphiopus holbrookii (Eastern Spadefoot) metamorphs were captured in the upland arrays, which lead us to estimate their abundance rather than having an exact count of individuals. For species with more than 15 captured individuals, we determined if they were more frequently found in upland 650 Southeastern Naturalist Vol. 10, No. 4 or lowland sites. We used the number of array days in each habitat to quantify sampling effort and conducted chi-square tests for goodness of fit, in which the expected values were based upon the proportion of total array days in the upland arrays (0.244) and the lowland arrays (0.756). Hog Island visual censuses From February 1992 to December 2001, we conducted 700 visual surveys of a 10-hectare mesic hammock that was surrounded by freshwater marsh on three sides and a shallow impoundment to the north (Fig. 1). These censuses were part of a long-term study of the site’s dense Sistrurus miliarius population (Farrell et al. 2008, May et al. 1996). Typically, 2 to 6 observers (mean = 4.2 people) visually searched the habitat for 2–3 hours (mean = 2.2 h) along a wide standard route that covered the study site and counted all herpetofauna (excluding chelonians). Chelonians, with the exception of Terrapene carolina (Florida Box Turtle) (Pilgrim et al. 1997), were rarely encountered and not counted. We identified members of three taxa (Hyla, Lithobates, and Plestiodon) to genera, since several congeners with similar appearance could be found in this study site. We did not turn cover objects during searches. Hog Island was dominated by Quercus virginiana Mill. (Live Oak), Sabal palmetto (Walter) Lodd. Ex Schultes & Schulthes (Cabbage Palm), Pinus elliottii Engelm. (Slash Pine), and Acer rubrum L. (Red Maple). We calculated the activity of each taxon by dividing the total number of individuals observed in a census by the number of observer hours involved in that census. Aquatic habitat sampling and terrestrial coverboard arrays We used visual surveys, auditory surveys, dip netting, and funnel traps to investigate the species richness of the aquatic herpetofauna. The funnel traps were designed as in Fitch (1987). Our efforts strongly focused on two impoundments (pool 1 and pool 3), with dip netting efforts conducted at least 4 times a year in each year from 1990 to 2010. We also had several hundred funnel-trap nights of effort from 2006 to 2010 in these impoundments. We used dip netting and funnel trapping in the cypress swamp habitat east of pool 1 and in the floating vegetation mats found along the southern edge of Spring Garden Lake. We also sampled 2 ephemeral ponds in the Volusia tract of LWNWR (USFWS 2008) using funnel traps from August to October in 2004. We placed 56 plywood coverboards (approximately 1-m x 1-m) in a variety of terrestrial habitats in 1995. These boards were checked weekly for 2 years and less frequently for the next three years. We recorded the species observed under the boards. Crepuscular levee censuses From April 1997 to November 1999, we conducted 110 visual censuses along the southern-most levee in the refuge. This 1600-m levee was bordered primarily by freshwater marsh and mesic hammock to the south and by 2 anthropogenic impoundments (pool 1 and pool 3) to the north (Fig. 1). Each census involved traveling west along the levee and then making the return trip (the total distance covered per census was 3200-m). We conducted at least 3 censuses each month, with an average of 7 censuses per month in the study period. We started each census no earlier than 30 minutes before sunset and ended approximately 90 minutes after sunset; we used 2011 T.M. Farrell, M.A. Pilgrim, P.G. May, and W.B. Blihovde 651 a head-lamp after darkness fell. We counted all reptiles and amphibians seen on the levee to develop quantitative estimates of species richness and activity. Results Refuge species richness We observed a total of 22 amphibian species and 50 reptile species on LWNWR (Appendix 1). The presence of 2 species, Rhineura floridana (Florida Wormlizard) and Crotalus horridus (Timber Rattlesnake), was documented Table 1. Herpetofauna captured in upland and lowland drift fence arrays. n = number of captures. % = the percentage of total snake, lizard, or anuran captures. Authorities and common names are listed in Appendix 1. *Scaphiopus not included in total upland counts for anurans. Upland sites Lowland sites Species n % Activity n % Activity χ2 P Snakes Agkistrodon piscivorus 0 0.0 0.0000 1 0.9 0.0008 Coluber constrictor 8 42.1 0.0203 42 41.2 0.0344 1.91 0.17 Micrurus fulvius 1 5.3 0.0025 0 0.0 0.0000 Nerodia fasciata 0 0.0 0.0000 7 6.9 0.0057 Pantherophis alleghaniensis 2 10.5 0.0051 10 9.8 0.0082 3.09 0.08 Pantherophis guttatus 0 0.0 0.0000 2 2.0 0.0016 Sistrurus miliarius 6 31.6 0.0152 15 14.7 0.0123 0.21 0.65 Thamnophis sauritus 1 5.3 0.0025 9 8.8 0.0074 Thamnophis sirtalis 1 5.3 0.0025 16 15.7 0.0131 Total snakes 19 100.0 0.0481 102 100.0 0.0835 Lizards Anolis carolinensis 17 28.3 0.0431 30 30.9 0.0246 3.48 0.06 Anolis sagrei 2 3.3 0.0051 4 4.1 0.0033 Aspidoscelis sexlineata 2 3.3 0.0051 0 0.0 0.0000 Plestiodon fasciatus 0 0.0 0.0000 2 2.1 0.0016 Plestiodon inexpectatus 11 18.3 0.0279 22 22.7 0.0180 1.38 0.24 Plestiodon laticeps 12 20.0 0.0305 6 6.2 0.0049 17.40 less than 0.01 Ophisaurus ventralis 0 0.0 0.0000 1 1.0 0.0008 Sceloporus undulatus 3 5.0 0.0076 0 0.0 0.0000 Scincella lateralis 13 21.7 0.0330 32 33.0 0.0262 0.48 0.49 Total lizards 60 100.0 0.1523 97 100.0 0.0795 Anurans Anaxyrus terrestris 2 5.3 0.0051 0 0.0 0.0000 Eleutherodactylus planirosterus 1 2.6 0.0025 0 0.0 0.0000 Gastrophryne carolinensis 16 42.1 0.0406 166 36.4 0.1361 24.00 less than 0.01 Hyla cinerea 0 0.0 0.0000 47 10.3 0.0385 15.20 less than 0.01 Hyla femoralis 6 15.8 0.0152 22 4.8 0.0180 0.12 0.72 Hyla squirella 1 2.6 0.0025 26 5.7 0.0213 6.29 0.01 Pseudacris ocularis 0 0.0 0.0000 9 2.0 0.0074 Lithobates clamitans 1 2.6 0.0025 6 1.3 0.0049 Lithobates sphenocephalus 11 28.9 0.0279 172 37.7 0.1410 33.60 <0.01 Lithobates grylio 0 0.0 0.0000 2 0.4 0.0016 Scaphiopus holbrookii* 300 na na 6 1.3 0.0049 898.00 <0.01 Total anurans 38 100.0 0.0964 456 100 0.3738 652 Southeastern Naturalist Vol. 10, No. 4 solely by photographs taken by LWNWR personnel, while the other 70 species were encountered during our sampling efforts. The amphibian fauna was dominated by anurans, as we observed only 5 species of salamanders. The reptile fauna was dominated by snake taxa, as 26 of the 50 reptile species we observed were snakes. Seven percent (5 of 72 species) of all species were introduced, and we found these non-native species primarily in disturbed habitats (Appendix 1). Drift fence arrays We captured a total of 29 species in the arrays (Table 1). The number of species captured in the upland arrays (21 species) was similar to the number captured in the lowland arrays (24 species), despite the larger number of arrays in lowland habitats. The Shannon-Wiener diversity index was higher for herpetofauna captured in lowland arrays (H' = 2.35) relative to upland arrays (H' = 1.32). The difference was largely a result of uneven species abundances in the upland captures, where S. holbrookii constituted 72% of all captures. Two species (Plestiodon laticeps [Broad-headed Skink] and S. holbrookii ) were significantly more abundant in upland areas than in the lowland areas, while 4 anuran species (Gastrophryne carolinensis [Eastern Narrow-mouthed Toad], Hyla cinerea [Green Treefrog], Hyla squirella [Squirrel Treefrog], and Lithobates sphenocephalus [Southern Leopard Frog]) were significantly more abundant in the lowland areas (Table 1). Seven common species (Coluber constrictor [Southern Black Racer], Pantherophis alleghaniensis [Eastern Ratsnake], Sistrurus miliarius, Anolis carolinensis [Southern Green Anole], Plestiodon inexpectatus [Southeastern Five-lined Skink], Scincella lateralis [Little Brown Skink], and Hyla femoralis [Pine Woods Treefrog]) were more evenly distributed with no statistically signifi- cant differences in capture rates between upland and lowland areas. Table 2. Snakes and anurans encountered during the crepuscular levee censuses. Ind./obs. hr. = number of individuals captured per observer hour. Species Total observed % of snakes Ind./obs. hr. Snakes Diadophis punctatus 2 0.7 0.013 Nerodia fasciata 132 45.5 0.885 Nerodia floridana 7 2.4 0.047 Nerodia taxispilota 2 0.7 0.013 Pantherophis alleghaniensis 4 1.4 0.027 Pantherophis guttatus 1 0.3 0.007 Regina alleni 1 0.3 0.007 Seminatrix pygaea 11 3.8 0.074 Sistrurus miliarius 27 9.3 0.181 Thamnophis sauritus 30 10.3 0.201 Thamnophis sirtalis 1 0.3 0.007 Totals for snakes 218 100.0 1.461 Anurans % of anurans Anaxyrus terrestris 5 1.7 0.034 Hyla cinerea 5 1.7 0.034 Lithobates grylio 11 3.8 0.074 Lithobates sphenocephalus 269 92.8 1.803 Totals for anurans 290 100.0 1.944 2011 T.M. Farrell, M.A. Pilgrim, P.G. May, and W.B. Blihovde 653 Crepuscular Levee Censuses We encountered only snakes and anurans during the levee censuses (Table 2). We observed 11 species of snakes. The counts were dominated by Nerodia fasciata (Florida Watersnake), Sistrurus miliarius, and Thamnophis sauritus (Peninsula Ribbonsnake), which together accounted for 86.7% of all observed snakes. The anuran sample was dominated by Lithobates sphenocephalus (Florida Leopard Frog), as this species accounted for 92.8% of all observed anurans (Table 2). Hog Island censuses We observed a total of 21 species (excluding chelonians) in this sampling effort, but 3 taxa (S. miliarius, Lithobates spp., and A. carolinensis) dominated the herpetofauna in terms of relative abundance (Table 3). Discussion The herpetofauna of LWNWR is highly diverse, with 72 species encountered during our study. Tuberville et al. (2005) surveyed the herpetofauna of 16 conservation areas maintained by the National Park Service in the Southeastern United States. None of these of these refuges had more than 64 documented native Table 3. Herpetofauna observed in visual censuses of Hog Island. n = the number of captures. Sampling effort is represented by individuals captured per observer hour (Ind./obs. hr). Species n % of snakes Ind./obs. hr Snakes Agkistrodon piscivorus 34 0.56 0.01 Coluber constrictor 80 1.32 0.02 Diadophis punctatus 3 0.05 <0.01 Lampropeltis triangulum 39 0.65 0.01 Micrurus fulvius 2 0.03 <0.01 Nerodia fasciata 43 0.71 0.01 Opheodrys aestivus 10 0.31 <0.01 Pantherophis alleghaniensis 13 0.21 <0.01 Seminatrix pygaea 1 0.02 <0.01 Sistrurus miliarius 5529 91.43 1.35 Thamnophis sauritus 222 3.67 0.05 Thamnophis sirtalis 61 1.00 0.02 Totals for snakes 6047 100.00 1.48 Lizards % of lizards Anolis carolinensis 1170 76.42 0.29 Plestiodon spp. 212 13.85 0.05 Ophisaurus ventralis 14 0.91 <0.01 Scincella lateralis 135 8.82 0.03 Totals for lizards 1531 100.00 0.38 Anurans % of anurans Anaxyrus terrestris 15 0.15 <0.01 Gastrophryne carolinensis 70 0.72 0.02 Hyla spp. 513 5.38 0.13 Pseudacris ocularis 52 0.54 0.01 Lithobates spp. 9065 93.31 2.21 Totals for anurans 9715 100.00 2.37 654 Southeastern Naturalist Vol. 10, No. 4 species of reptiles and amphibians. With 67 documented native species, LWNWR exhibits great species richness; the total number of species captured represents 82.9% of Volusia County Florida’s 82 native, non-marine species (Ashton and Ashton 1981, 1985, 1988; and T.M. Farrell, pers. observ.). Tuberville et al. (2005) found that park area was strongly associated with species richness (log Richness = 0.274 * log area + 0.722) in the Southeast Park Network. Using this function, LWNWR is expected to contain 63.3 native species, which is approximately 4 fewer species than the observed total. Thus, LWNWR has high richness even after adjusting for the refuge’s large area. We attribute this richness to the high diversity of both terrestrial and aquatic habitats on the refuge, and to the proximity of the refuge to vast tracts of protected land, including Ocala National Forest, DeLeon Springs State Park, and Lake George State Forest. The richness data we presented are likely to provide a fairly accurate assessment of the current herpetofauna in the more accessible portions of the refuge, given the diversity of sampling methods used and the length of time and large number of observer hours involved in our efforts. Ryan et al. (2002) found that the diversity of sampling methods employed in a survey is likely to increase documented species richness. One indication of the completeness of our inventory is that we have only documented the presence of 2 additional species (Crotalus horridus [Timber Rattlesnake] and Notophthalmus viridescens [Peninsula Newt]) during the last 5 years of fieldwork. For C. horridus, we have a single record (Blihovde and Brady 2010) and wonder if this animal was moved by humans or represents a true southern range extension for this species. The herpetofauna of peninsular Florida is undergoing major changes, with large declines of some native species (e.g., Krysko and Smith 2005) and a massive influx of introduced species (Meshaka et al. 2004). Continued monitoring of the diverse herpetofauna of Lake Woodruff National Wildlife Refuge could facilitate a better understanding of factors underlying ongoing changes in species abundances and community assemblages that are currently occurring in the ecosystems of peninsular Florida. Acknowledgments We extend our gratitude to more than 60 Stetson University undergraduates who helped conduct field surveys. We had major support from the Cici and Hyatt Brown Faculty Fellow Program at Stetson. We also thank the current and former staff of Lake Woodruff National Wildlife Refuge, especially Leon Rhodes, Harold Morrow, Kristina Sorenson, and Brian Braudis. Literature Cited Ashton, R.E., and P.S. Ashton. 1981. Handbook of Reptiles and Amphibians of Florida, Part One: The Snakes. Windward Publishing, Inc., Miami, FL. 176 pp. Ashton, R.E., and P.S. Ashton. 1985. Handbook of Reptiles and Amphibians of Florida, Part Two: Lizards, Turtles and Crocodilians. Windward Publishing, Inc., Miami, FL. 191 pp. Ashton, R.E., and P.S. Ashton. 1988. Handbook of Reptiles and Amphibians of Florida, Part Three: The Amphibians. Windward Publishing, Inc., Miami, FL. 191 pp. 2011 T.M. Farrell, M.A. Pilgrim, P.G. May, and W.B. Blihovde 655 Beaupre, S.J., and L.E. Douglas. 2009. Snakes as indicators and monitors of ecosystem properties. Pp. 244–261, In S.J. Mullin and R.A. Seigel (Eds.). Snakes: Ecology and Conservation. Cornell University Press, Ithaca, NY. 365 pp. Blihovde, W.B., and K.A. Brady. 2010. Crotalus horridus-Geographic distribution. Herpetological Review 41:244. Crother, B.I. (Ed.). 2008. Scientific and standard English names of amphibians and reptiles of North America North of Mexico. SSAR Herpetological Circular 37. 84 pp. Davic, R.D., and H.H. Welsh, Jr. 2004. On the ecological role of salamanders. Annual Review of Ecology, Evolution, and Systematics 35:405–434. Dial, R., and J. Roughgarden. 1995. Experimental removal of insectivores from rain forest canopy: Direct and indirect effects. Ecology 76:1821–1834. Farrell, T.M., M.A. Pilgrim, and P.G. May. 2008. Annual variation in neonate recruitment in a Florida population of the Pigmy Rattlesnake (Sistrurus miliarius). Pp. 257–265, In W. Hayes, M. Cardwell, S. Bush, and K. Beaman (Eds.). The Biology of Rattlesnakes. Loma Linda Press, Loma Linda, CA. 606 pp. Fitch, H.S. 1987. Collecting and life-history techniques. Pp. 143–164, In R.A. Seigel, J.T. Collins, and S.S. Novak (Eds.). Snakes: Ecology and Evolutionary Biology. Macmillan, New York, NY. 529 pp. Gibbons, J.W., D.E. Scott, T.J. Ryan, K.A. Buhlmann, T.D. Turberville, B.S. Metts, J.L. Greene, T. Mills, Y. Leiden, S. Poppy, and C.T. Winne. 2000. The global decline of reptiles: Déjà vu amphibians. Bioscience 50:653–666. Houlahan, J.E., C.S. Findlay, B.R. Schmidt, A.H. Meyer, and S.L. Kuzmin. 2000. Quantitative evidence for global amphibian population declines. Nature 404:752–755. Krysko, K.L., and D.J. Smith. 2005. The decline and extirpation of the Kingsnake in Florida. Pp. 132–141, In W. E. Meshaka and K. J. Babbitt (Eds.). Status and Conservation of Florida Amphibians and Reptiles. Krieger Press, Malabar, FL. 334 pp. May, P.G., T.M. Farrell, S.T. Heulett, M.A. Pilgrim, L.A. Bishop, D.J. Spence, A.M. Rabatski, M.G. Campbell, A.D Aycrigg, and W.E. Richardson. 1996. The seasonal abundance and activity of a rattlesnake (Sistrurus miliarius barbouri) in central Florida. Copeia 1996:389–400. Meshaka, W.E., B.P. Butterfield, and J.B. Hauge. 2004. The Exotic Amphibian and Reptiles of Florida. Krieger Press, Malabar, FL. 155 pp. Pilgrim, M.A., T.M. Farrell, and P.G. May. 1997. Population structure, activity, and sexual dimorphism in a central Florida population of Terrapene carolina. Chelonian Conservation and Biology 2:483–488. Regester, K.J., K.R. Lips, and M.R. Whiles. 2006. Energy flow and subsidies associated with the complex life cycle of ambystomatid salamanders in ponds and adjacent forest in southern Illinois. Oecologia 147:303–314. Ryan, T.J., T. Philippi, Y.A. Leiden, M.E. Dorcas, T.B. Wigley, and J.W. Gibbons. 2002. Monitoring herpetofauna in a managed forest landscape: Effects of habitat types and census techniques. Forestry Ecology and Management 167:83–90. Tuberville, T.D., J.D. Willson, M.E. Dorcas, and J.W. Gibbons. 2005. Herpetofaunal diversity of southeastern National Parks. Southeastern Naturalist 4:537–569. US Fish and Wildlife Service (USFWS). 2008. Lake Woodruff National Wildlife Refuge: Comprehensive Conservation Plan. Available online at planning/PDFdocuments/ LakeWoodruffFinalCCP/Final%20CCP%20Lake%20 Woodruff.pdf. Accessed 15 July 2010. Welsh, H.H., Jr., and S. Droege. 2001. A case for using plethodontid salamanders for monitoring biodiversity and ecosystem integrity of North American forests. Conservation Biology 15:558–568. 656 Southeastern Naturalist Vol. 10, No. 4 Appendix 1. Amphibian and reptile species found between 1990 and 2010 at LWNWR (* indicates introduced species). Habitat codes indicate the habitat where we most often encountered each species: xh = xeric hammock, mh = mesic hammock, pf = pine flatwoods, cs = cypress swamp, fm = freshwater marsh, tp = temporary pools, pw = permanent waters, da = human disturbed areas. The taxonomy follows Crother (2008). Latin name Common name xh mh pf cs fm tp pw da Anura: Frogs and toads Acris gryllus (LeConte, 1825) Florida Cricket Frog X Anaxyrus quercicus (Holbrook, 1840) Oak Toad X X Anaxyrus terrestris (Bonnaterre, 1789) Southern Toad X X X X Eleutherodactylus planirostris* (Cope, 1862) Greenhouse Frog X X X Gastrophryne carolinensis (Holbrook, 1836) Eastern Narrow-mouthed Toad X X X Hyla cinerea (Schneider, 1799) Green Treefrog X X X X Hyla femoralis Bosc, 1800 Pine Woods Treefrog X X X Hyla squirella Bosc, 1800 Squirrel Treefrog X X X X Lithobates clamitans (Latreille, 1801) Bronze Frog X X Lithobates grylio (Stejneger, 1901) Pig Frog X Lithobates heckscheri (Wright, 1924) River Frog X Lithobates sphenocephalus (Cope, 1886) Florida Leopard Frog X X X X X Osteopilus septentrionalis* (Dumeril and Bibron, 1841) Cuban Treefrog X X Pseudacris crucifer (Wied Neuwied, 1838) Spring Peeper X Pseudacris nigrita (Le Conte, 1825) Southern Chorus Frog X Pseudacris ocularis (Bosc and Daudin, 1801) Little Grass Frog X Scaphiopus holbrookii (Harlan, 1835) Eastern Spadefoot X Caudata: Salamanders Amphiuma means Garden, 1821 Two-toed Amphiuma X Eurycea quadridigitata (Holbrook, 1842) Dwarf Salamander X Notophthalmus viridescens (Rafinesque, 1820) Peninsula Newt X Plethodon grobmani Allen and Neill, 1949 Southeastern Slimy Salamander X X Siren lacertina L., 1766 Greater Siren X 2011 T.M. Farrell, M.A. Pilgrim, P.G. May, and W.B. Blihovde 657 Latin name Common name xh mh pf cs fm tp pw da Squamata: Lizards and amphisbaenians Anolis carolinensis (Voight, 1832) Southern Green Anole X X X X X X Anolis sagrei* Dumeril and Bibron, 1837 Cuban Brown Anole X Aspidoscelis sexlineata (L., 1766) Eastern Six-lined Racerunner X Hemidactylus garnotii* Dumeril and Bibron, 1836 Indo-Pacific House Gecko X Ophisaurus ventralis (Linnaeus, 1766) Eastern Glass Lizard X X X X Plestiodon egregius (Baird, 1859) Peninsula Mole Skink X Plestiodon fasciatus (L., 1758) Common Five-lined Skink X X X Plestiodon inexpectatus Taylor, 1932 Southeastern Five-lined Skink X X X Plestiodon laticeps (Scheider, 1801) Broad-headed Skink X X X Rhineura floridana (Baird, 1859) Florida Wormlizard X Sceloporus undulatus (Bosc and Daudin in Sonnini Eastern Fence Lizard X and Latreille, 1801) Scincella lateralis (Say in James, 1832) Little Brown Skink X X X Squamata: Snakes Agkistrodon piscivorus (Lacepede, 1789) Florida Cottonmouth X X X Cemophora coccinea (Blumenbach, 1788) Florida Scarletsnake X Coluber constrictor L., 1758 Southern Black Racer X X X X X Coluber flagellum Shaw, 1802 Eastern Coachwhip X Crotalus adamanteus Palisot de Beauvois, 1799 Eastern Diamond-backed Rattlesnake X X X Crotalus horridus Linnaeus, 1758 Timber Rattlesnake X Diadophis punctatus (Linnaeus, 1766) Southern Ring-necked Snake X X X Farancia abacura (Holbrook, 1836) Eastern Mudsnake X X Lampropeltis triangulum (Lacepede, 1789) Scarlet Kingsnake X X Micrurus fulvius (L., 1766) Harlequin Coralsnake X X Nerodia fasciata (L., 1766) Florida Watersnake X X X Nerodia floridana (Goff, 1936) Florida Green Watersnake X Nerodia taxispilota (Holbrook, 1838) Brown Watersnake X Opheodrys aestivus (L., 1766) Florida Rough Green Snake X X 658 Southeastern Naturalist Vol. 10, No. 4 Latin name Common name xh mh pf cs fm tp pw da Pantherophis alleghaniensis (Holbrook, 1836) Eastern Ratsnake X X X Pantherophis guttatus (L., 1766) Red Cornsnake X X X Pituophis melanoleucus (Daudin, 1803) Florida Pinesnake X Regina alleni (Garman, 1874) Striped Crayfish Snake X Rhadinaea flavilata (Cope, 1871) Pine Woods Littersnake X Seminatrix pygaea (Cope, 1871) Black Swampsnake X Sistrurus miliarius (L., 1766) Dusky Pigmy Rattlesnake X X X X Storeria dekayi (Holbrook, 1836) Midland Brownsnake X Storeria occipitomaculata (Storer, 1839) Florida Red-bellied Snake X Tantilla relicta Telford, 1966 Central Florida Crowned Snake X Thamnophis sauritus (L., 1766) Peninsula Ribbonsnake X X X Thamnophis sirtalis (L., 1758) Eastern Gartersnake X X X X Crocodilians: Alligators Alligator mississipiensis (Daudin, 1801) American Alligator X Testudines: Turtles and tortoises Apalone ferox (Schneider, 1783) Florida Softshell X Chelydra serpentina (L., 1758) Florida Snapping Turtle X X Deirochelys reticularia (Latreille, 1801) Florida Chicken Turtle X Gopherus polyphemus (Daudin, 1802) Gopher Tortoise X Kinosternon baurii (Garman, 1891) Striped Mud Turtle X X Pseudemys nelsoni Carr, 1938 Florida Red-bellied Turtle X Pseudemys peninsularis Carr, 1938 Peninsula Cooter X Sternotherus minor (Agassiz, 1857) Loggerhead Musk Turtle X Sternotherus odoratus (Latreille, 1801) Eastern Musk Turtle X Terrapene carolina (L., 1758) Florida Box Turtle X X X X Trachemys scripta* (Schoepf, 1792) Red-eared Slider X