Southeastern Naturalist J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 182 2017 SOUTHEASTERN NATURALIST 16(2):182–194 Using Drift Fence Arrays to Compare Terrestrial Herpetofauna Diversity in Three Habitats at Tishomingo National Wildlife Refuge, Oklahoma John A. Muller1,*, Joseph A. Veech1, and Justin Roach2 Abstract- Unlike avifauna, which have been inventoried and monitored for decades at Tishomingo National Wildlife Refuge, OK, herpetofauna have had few targeted inventories on the refuge. Therefore, in 2010 we established 6 permanent drift fence arrays to expand the current knowledge on the terrestrial herpetofauna present within the refuge. The arrays were in operation for a total of 105 nights over 5 years (2010–2014), capturing 1122 vertebrates including 12 anuran, 1 salamander, 13 snake, 5 lizard, and 2 turtle species. The surveys resulted in the discovery of 5 new amphibian species as well as a new mammal species on the refuge. By placing arrays across 3 different habitat types, we were able to compare and determine that there were differences in species diversity and relative abundance across those habitats. The results have shown the richness of herpetofauna in the area has been underestimated. The study also indicates that a preserve established for the conservation of one particular species group (waterfowl and neotropical migratory birds) can have appreciable diversity of another vertebrate group. Introduction Loss of biodiversity has become one of the leading concerns in conservation science. Wilson (1988) set the stage for the current view of the state of global biodiversity, including concerns over the rapid rate of species loss and the consequences that those losses entail (Ceballos et al. 2015). Amphibians are one of the fastest declining vertebrate groups worldwide. Recently, the International Union for the Conservation of Nature (IUCN 2008) listed 32.4% of all known amphibian species as threatened or extinct. Declines in amphibian species have been attributed to disease, habitat destruction, environmental contaminants, and global climate change (Alford and Richards 1999, Collins and Storfer 2003). Because they are sensitive to a variety of environmental factors, both reptiles (Beaupre and Douglas 2009) and amphibians (Welsh and Droege 2001) are considered bioindicators of ecosystem health and should be deemed important groups for conservation. Amphibians are especially useful in areas where toxins are present (such as near agricultural operations) because of their sensitivity to chemicals, even at very small levels in the environment (Hayes et al. 2006). Although not currently considered polluted in any way, Tishomingo National Wildlife Refuge (NWR) in Johnston County, OK, was formerly an area of very intensive agriculture, and agriculture is still 1Department of Biology, Texas State University, 601 University Drive, San Marcos, TX 78666. 2Tishomingo National Wildlife Refuge, 12000 S Refuge Road, Tishomingo, TX73460. *Corresponding author - jamuller@austin.rr.com. Manuscript Editor: Brad Glorioso Southeastern Naturalist 183 J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 very prominent in surrounding areas (USFWS 2010). As conventional agricultural practices rely on the use of pesticides, herbicides, fertilizers, and other chemicals, continued amphibian monitoring could be used to indicate the potential presence of pollutants harmful to the natural community at Tishomingo NWR. Management actions on public lands can directly affect species abundance (Petranka et al. 1993, Renken et al. 2004). Therefore, managers of public lands should consider it a priority to fully determine the current status of their biodiversity before taking management actions that might inadvertently impact non-focal species for which those lands also provide important habitat. Tishomingo NWR was established in 1946 for the management and research of migratory waterfowl. Currently, the refuge still manages for waterfowl through farming programs and wetland management. Recently, Tishomingo NWR has participated in more research and management of various other taxonomic groups, including Protonotaria citrea (Boddaert) (Prothonotary Warbler), Alnus maritima (Marshall) Muhl. ex Nutt. (Seaside Alder), and Macrochelys temminckii (Troost in Harlan) (Alligator Snapping Turtle), making the overall conservation mission of the refuge more comprehensive. Inventory and monitoring is a useful tool for tracking current levels of and changes to biodiversity. Complete surveys of herpetofaunal communities are nonexistent for many areas, including some protected and conservation areas. Therefore, establishing complete species lists and relative abundances of species is a priority for many such areas, including Tishomingo NWR. As of 2010, there were 65 species of herpetofauna on the species list at Tishomingo NWR (USFWS 2010). This list was derived from a combination of spontaneous staff and volunteer sightings, publications (e.g., Carpenter 1958), specimen collections such as the one found at University of Oklahoma’s Sam Noble Oklahoma Museum of Natural History, as well as multiple species that occur in the area that have not yet been seen on the refuge. These sources, however, do not give relative abundances and some sources are from the surrounding area and not the refuge itself. Specifically, the goal of our monitoring program was to confirm species that occur within the refuge and determine their relative abundances to provide important baseline data for future monitoring efforts. To this end, we conducted a 5-year terrestrial herpetological inventory of Tishomingo NWR employing drift-fence arrays to detect herpetofauna because they allow for long-term monitoring and are relatively simple to implement. Field-site Description Tishomingo National Wildlife Refuge is located in south-central Oklahoma along the northern extent of Lake Texoma (34º11.72'N 96º38.24'W; Fig. 1). Before the refuge was established, the land was part of the ~12,138-ha Washita Farm, which was known as the showcase of agriculture for southern Oklahoma. In 1944, after the construction of Denison Dam and the creation of Lake Texoma, the property was transferred to the US Army Corps of Engineers (USACE) under the Flood Control Act of 28 June 1938 (Public Law No. 761, 75th Congress, 3rd Session). The refuge is 6661 ha, of which ~2427 ha are encompassed by Cumberland Pool. Habitats within the refuge include upland forest, bottomland hardwood forest, riparian Southeastern Naturalist J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 184 areas, grasslands, and agricultural fields. Of the available habitats, we sampled grasslands, bottomland hardwoods, and upland forest. Methods Drift-fence arrays We placed 2 permanent drift-fence arrays in separate patches in each of the 3 habitat types. Each array was opened and checked back-to-back every day for 1 week each month when weather permitted herpetofauna to be active (usually March through September). To prevent bias from weather and other variables that could affect catch rate, all 6 arrays were open simultaneously and checked as close together as possible. Though drift-fence arrays can be arranged in several different configurations (Corn 1994, Gibbons and Semlitsch 1982, Hanlin et al. 2000), we designed our arrays in the shape of an X with each wing pointing toward one of the 4 cardinal directions. Each wing was 10 m long and made of 1-m-high sheet metal buried ~8 cm into the ground. We used 19-L plastic buckets as pitfalls placed at the end of each wing, with 1 also placed at the intersection of all 4 wings. We placed a double-sided funnel trap in the center along both sides of each wing. Funnel traps were made of 0.6-cm hardware cloth and were 1 m in length and 0.2 m in diameter. We placed wet sponges in each bucket to reduce mortality due to desiccation. Figure 1. Map showing the location of Tishomingo National Wildlife Refuge, with an inset showing the location of the refuge in relation to the state of Oklahoma. Southeastern Naturalist 185 J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 Upland and bottomland sites were prone to flooding, so we drilled holes into the bottom of each bucket allowing water to flow out and placed small blocks of wood as “floaters” for captured individuals to climb onto to stay above water. We used cover boards to provide shade for each pitfall and funnel. Other than checking traps daily, we made no attempt to prevent predation (e.g., by mesocarnivores) on trap occupants. Some predation may have occurred, although we did not see obvious signs (e.g., tracks) of predator visitation. We used various field guides to identify captured individuals (e.g., Conant and Collins 1998, Sievert and Sievert 2005). We also took photographs for species identification and documentation as well as for future reference. Species diversity To compare species diversity across habitats, we used the Shannon index (H'; Krebs 1999). However, because this index is influenced by species richness in which habitats with higher species richness have greater potential H', we converted H' to evenness (J'; Zar 1999). J' represents how close the community is to being evenly distributed, or how close the community is to being maximally diverse. Species abundance Differences in species abundance among habitat types were tested using the chisquare goodness-of-fit test applied to each species separately. Relative abundance of individual species was compared, pairwise, across habitats (bottomland forest vs. upland forest, bottomland forest vs. grassland, upland forest vs. grassland). We conducted the pairwise comparisons, instead of a chi-square test applied simultaneously to all 3 habitat types, to determine which exact habitats might differ from one another rather than whether there were differences overall. Because all 6 arrays were run simultaneously and for an equal amount of time, trap effort and potential capture were assumed to be even across habitats. Because some species occurred in low frequencies, we included only species with expected frequencies of greater than 5 (Sokal and Rohlf 1995). We controlled for false discovery rate (study-wide Type I error rate) by adjusting the critical value as in Benjamini and Yekutieli (2001); for 20 comparisons and an original α = 0.05, the adjusted α = 0.014. Results We opened and checked the 6 arrays daily for a total of 27 weeks over 5 years (2010–2014). There were a total of 1122 captures composed of 33 species (12 anuran, 1 salamander, 13 snake, 5 lizard, and 2 turtle; Table 1). Two individual frogs eaten by fire ants were unidentifiable and excluded from analyses . Among the 3 habitat types, bottomland forest had the highest number of amphibians captured (485) but the lowest number of reptiles (28). Grasslands had the highest number of reptiles captured (243) but the lowest number of amphibians (126). Upland forest had 54 reptiles and 186 amphibians captured. Bottomland forest and upland forest captures were dominated by a single species, Lithobates sphenocephalus (Cope) (Southern Leopard Frog), with 65% and 52% of captures, Southeastern Naturalist J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 186 Table 1. Species captured with the number per habitat and percentage that each species constituted from each habitat type. Bottomland Upland Grassland Species with common name Captures % Captures % Captures % Anurans Acris blanchardi Harper 7 1.30% 1 0.39% 1 0.25% (Blanchard’s Cricket Frog) Anaxyrus americanus 45 8.36% 8 3.15% 2 0.49% (American Toad) Anaxyrus woodhousii 0 0.00% 3 1.18% 1 0.25% (Woodhouse's Toad) Gastrophryne carolinensis 51 9.48% 30 11.81% 6 1.48% (Eastern Narrow-mouthed Toad) Gastrophryne olivacea 1 0.19% 1 0.39% 2 0.49% (Western Narrow-mouthed Toad) Hyla cinerea 1 0.19% 1 0.39% 1 0.25% (Green Treefrog) Hyla versicolor/chrysoscelis 7 1.30% 0 0.00% 1 0.25% (Gray Treefrog Complex) Lithobates catesbeianus 1 0.19% 0 0.00% 9 2.22% (American Bullfrog) Lithobates palustris 17 3.16% 1 0.39% 2 0.49% (Pickerel Frog) Lithobates sphenocephalus 348 64.68% 131 51.57% 100 24.69% (Southern Leopard Frog) Pseudacris fouquettei Moriarty Lemmon, 3 0.56% 4 1.57% 0 0.00% Lemmon, Collins and Cannatella (Cajun Chorus Frog) Scaphiopus hurterii 0 0.00% 3 1.18% 0 0.00% (Hurter’s Spadefoot) Lizards Cnemidophorus sexlineatus 0 0.00% 21 8.27% 209 51.60% (Six-lined Racerunner) Plestiodon fasciatus (L.) 9 1.67% 2 0.79% 0 0.00% (Common Five-lined Skink) Plestiodon laticeps (Schneider, 1801) 1 0.19% 3 1.18% 0 0.00% (Broad-headed Skink) Sceloporus consobrinus (Baird and Girard) 0 0.00% 8 3.15% 0 0.00% (Prairie lizard) Scincella lateralis (Mittleman) 3 0.56% 4 1.57% 0 0.00% (Little Brown Skink) Snakes Agkistrodon contortrix (L. 0 0.00% 3 1.18% 0 0.00% (Copperhead) Agkistrodon piscivorus (Lacepede) 1 0.19% 0 0.00% 3 0.74% (Cottonmouth) Coluber constrictor 0 0.00% 2 0.79% 11 2.72% (Racer) Crotalus horridus L. 0 0.00% 1 0.39% 0 0.00% (Timber Rattlesnake) Southeastern Naturalist 187 J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 respectively. Grassland was dominated by 2 species, Cnemidophorus sexlineatus (L.) (Six-Lined Racerunner) and Southern Leopard Frog with 52% and 25% of captures, respectively. Table 1, continued. Bottomland Upland Grassland Species with common name Captures % Captures % Captures % Diadophis punctatus Baird and Girard 0 0.00% 1 0.39% 0 0.00% (Ring-necked Snake) Haldea striatula Baird and Girard 1 0.19% 1 0.39% 0 0.00% (Rough Earthsnake) Lampropeltis calligaster (Harlan) 1 0.19% 1 0.39% 0 0.00% (Yellow-bellied Kingsnake) Pantherophis emoryi (Baird and Girard) 1 0.19% 0 0.00% 0 0.00% (Great Plains Ratsnake) Pantherophis obsoletus (Say in James) 1 0.19% 4 1.57% 5 1.23% (Western Ratsnake) Sistrurus miliarius (L.) 3 0.56% 3 1.18% 0 0.00% (Pygmy Rattlesnake) Storeria dekayi (Holbrook) 6 1.12% 0 0.00% 0 0.00% (Dekay’s Brownsnake) Thamnophis proximus 3 0.56% 2 0.79% 13 3.21% (Western Ribbonsnake) Virginia valeriae (Baird and Girard) 0 0.00% 1 0.39% 0 0.00% (Smooth Earthsnake) Salamanders Notophthalmus viridescens 1 0.19% 0 0.00% 1 0.25% (Eastern Newt) Turtles Terrapene Carolina (L.) 0 0.00% 0 0.00% 1 0.25% (Eastern Box Turtle) Trachemys scripta (Thunberg in Schoepff) 1 0.19% 0 0.00% 1 0.25% (Pond Slider) Mammals Blarina hylophaga 1 0.19% 0 0.00% 0 0.00% (Elliot’s Short-tailed Shrew) Cryptotis parva (Say) 6 1.12% 3 1.18% 11 2.72% (Least Shrew) Didelphis virginiana Kerr 1 0.19% 0 0.00% 0 0.00% (Virginia Opossum) Microtus pinetorum (Le Conte) 2 0.37% 0 0.00% 1 0.25% (Woodland Vole) Peromyscus spp. 5 0.93% 8 3.15% 5 1.23% (deer mouse spp.) Reithrodontomys fulvescens J. A. Allen 6 1.12% 3 1.18% 11 2.72% (Fulvous Harvest Mouse) Sciurus carolinensis Gmelin 2 0.37% 0 0.00% 0 0.00% (Eastern Gray Squirrel) Sigmodon hispidus Say and Ord 2 0.37% 0 0.00% 8 1.98% (Hispid Cotton Rat) Southeastern Naturalist J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 188 Upland forest had the highest overall diversity across all taxa (H' = 1.85, J' = 0.58) as well as the highest number of unique species (6) (Table 2). Bottomland forest had the highest number of captures (513) but had the lowest diversity (H' = 1.31, J' = 0.42). Grassland had the fewest unique species (1) and a diversity intermediate between that of upland and bottomland forests (H' = 1.35, J' = 0.47). Reptile diversity was greatest in the bottomland (H' = 2.13, J' = 0.86) and upland forests (H' = 2.18, J' = 0.81) whereas reptile diversity in grasslands was comparatively low (H’=0.61, J’=0.31). Amphibian diversity was fairly similar across all 3 habitats (Table 2). Of the 20 pairwise comparisons of species’ relative abundances between habitats, 13 were statistically significant (Table 3). These significant differences involved 7 species: Anaxyrus americanus (Holbrook) (American Toad), Sixlined Racerunner, Coluber constrictor L. (Racer), Gastrophryne carolinensis (Holbrook) (Eastern Narrow-mouthed Toad), Lithobates palustris (LeConte) (Pickerel Frog), Southern Leopard Frog, and Thamnophis proximus (Say in James) (Western Ribbonsnake). Of the 3 pairwise habitat comparisons, the greatest number of significant differences in relative abundances was between bottomland forest and grassland (6 species; Table 3). Six-lined Racerunner exhibited the strongest habitat differences; it was the only species to have significantly different relative abundances in all 3 habitat types (209, 21, and 0 in grassland, upland forest, and bottomland forest respectively; Table 3). The mostabundant and widely distributed species, the Southern Leopard Frog, was found in all 3 habitat types, although it was significantly more abundant in bottomland forest (348 captures) than in either upland forest or grassland (131 and 100 captures, respectively; Table 3). Table 2. Diversity indices for each combination of habitat type and taxon. H' = Shannon index, J' = evenness. # of Habitat # of captures # of species unique species H' J' Combined Bottomland 513 23 2 1.31 0.42 Upland 240 25 6 1.85 0.58 Grassland 369 18 1 1.35 0.47 All habitats 1122 33 1.76 0.50 Amphibians Bottomland 485 11 0 1.02 0.42 Upland 186 10 1 1.00 0.45 Grassland 126 11 0 0.91 0.38 All habitats 797 13 1.00 0.32 Reptiles Bottomland 28 12 2 2.13 0.86 Upland 54 15 5 2.18 0.81 Grassland 243 7 1 0.61 0.31 All habitats 325 20 1.37 0.45 Southeastern Naturalist 189 J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 Discussion Our results showed that the overall species richness of herpetofauna at Tishomingo NWR is greater than previously recorded, and given that only 3 potential habitats were surveyed, it is likely that the total number of species on the refuge is greater still. Among the 3 habitats that were surveyed, the species compositions varied in each, indicating that each habitat has a somewhat unique herpetofaunal community. Of the 13 species of amphibians captured, all but Anaxyrus woodhousii (Girard) (Woodhouse’s Toad) and Gastrophryne olivacea (Hallowell) (Western Narrowmouthed Toad) are eastern species. Four of the amphibians were found west of their expected range and were new to the refuge: Hyla cinerea (Schneider) (Green Treefrog), Notophthalmus viridescens (Rafinesque) (Eastern Newt), Pickerel Frog, and Eastern Narrow-mouthed Toad. These westward expansions of former ranges are similar to those of some mammals. Sciurus niger L. (Eastern Fox Squirrel) and Perimyotis subflavus (F. Cuvier) (Tri-colored Bat) are thought to be using riparian corridors to expand their ranges west (Geluso and Jones 2004, White et al. 2006). It is plausible these eastern amphibians are also using newly created riparian corridors to expand their range westward. Most of Tishomingo NWR’s forest was removed when it was an agricultural operation early in the 19th century, but since Table 3. Chi-square (χ2) values for the 20 species comparisons that had an expected frequency of at least 5. * indicate significance with false discovery rate adjustment of α = 0.014. BL = bottomland forest, UP = upland forest, and GR = Grassland. Species χ2 P Bottomland forest vs. upland forest BL UP American Toad 45 8 25.83 less than 0.001* Six-lined Racerunner 0 21 21.00 less than 0.001* Eastern Narrow-mouthed Toad 51 30 5.44 less than 0.025 Pickerel Frog 17 1 14.22 less than 0.005* Southern Leopard Frog 348 131 98.31 less than 0.001* American Five-lined Skink 9 2 4.45 less than 0.05 Upland forest vs. grassland UP GR American Toad 8 2 3.60 less than 0.1 Six-lined Racerunner 21 209 153.67 less than 0.001* Racer 2 11 6.23 less than 0.025 Eastern Narrow-mouthed Toad 30 6 16.00 less than 0.01* Southern Leopard Frog 131 100 4.16 less than 0.05 Western Ribbon Sanke 2 13 9.62 less than 0.001* Bottomland forest vs. grassland BL GR American Toad 45 2 39.34 less than 0.001* Six-lined Racerunner 0 209 209.00 less than 0.001* Racer 0 11 11.00 less than 0.001* Eastern Narrow-mouthed Toad 51 6 35.53 less than 0.001* American Bullfrog 1 9 6.40 less than 0.025 Pickerel Frog 17 2 11.84 less than 0.01* Southern Leopard Frog 348 100 137.29 less than 0.001* Western Ribbon Sanke 3 13 6.25 less than 0.025 Southeastern Naturalist J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 190 the area has been converted into a conservation area, the potential corridor forest has become reestablished. Other changes in the area have also helped to facilitate the expansion of these eastern forests, including the establishment of Denison Dam and the creation of Lake Texoma, which has created riparian habitats on the refuge (T. Patton, Southeastern Oklahoma State University, Durant, OK, 25 March 2015 pers. comm.). Bottomland forest is the dominant nonaquatic habitat type at Tishomingo NWR, encompassing about 2870 ha. It had the highest number of captures for the study (513) and a high species richness (23), although it had the lowest diversity index due to the dominance of Southern Leopard Frog captures (68%). Even though there is about 4.5 times less upland forest habitat than bottomland, upland forest had both the highest diversity index and the highest number of unique species (6). This indicates that although upland forest constitutes relatively little land area, it has great importance for multiple species and overall diversity within the refuge. The grasslands had the second most captures (369) and the second highest diversity index but the lowest species richness (18). The low species richness is likely due to the limited amount of grassland (161 ha). There may not be enough area in grasslands to sustain populations of some species, and hence they are absent from or temporally sporadic (due to extinction–recolonization) in the grassland habitat. Herpetofauna are typically abundant in ecosystems, and therefore they have important roles as both predator and prey (Davic and Welsh 2004, Dial and Roughgarden 1995, Regester et al. 2006). Consequently, any species captured in high numbers is likely an important member of the ecological community. Two species dominated all captures: Southern Leopard Frogs and Six-lined Racerunners constituted 52% and 20% of all captures, respectively. Both species likely serve as important predators upon invertebrates as well as common prey items for larger vertebrates in the community. Drift fences are an effective way to sample certain species of herpetofauna (Todd et al. 2007). They are particularly useful in determining the presence of cryptic species such as fossorial, or leaf-litter inhabitants, but they do have limitations. The results of this study most likely underestimate the herpetofaunal diversity of Tishomingo NWR. There are multiple species that occur frequently on the refuge but were not documented during this project. According to USFWS (2010), 65 species of reptiles and amphibians potentially occur on the refuge, meaning this survey only found 58% of potential species. One reason for this seemingly low percentage of overall species is that drift-fence arrays were located away from riparian habitats and open water, which are major habitat types on the refuge. This led to species that are commonly seen on the refuge, such as Nerodia spp. (watersnakes), being completely absent from the surveys. Some groups of species, such as larger snakes and turtles, are also harder to capture (because of their size) with pitfall and funnel traps (Enge and Marion 1986, Fitch 1992). Arboreal species such as Hyla versicolor LeConte (Gray Treefrog) and Hyla chrysoscelis Cope (Cope’s Gray Treefrog) are difficult to detect using terrestrial arrays (Gibbons and Semlitsch 1982). Large, fully aquatic herpetofauna such as Alligator Snapping Turtles would be difficult to detect Southeastern Naturalist 191 J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 with any terrestrial monitoring protocol. Yet even though there are limitations to using drift fences and pitfall traps as a singular method to determine species presence on the refuge, multiple new species including Green Treefrog, Eastern Newt, Pickerel Frog, Eastern Narrow-mouthed Toad, and Scaphiopus hurterii Strecker (Hurter’s Spadefoot) were detected using this sampling protocol. Even though the intention of the project was to investigate the diversity of herpetofauna on the refuge, the drift fence and pitfall arrays documented the presence of other taxonomic groups. At least 8 mammal species were captured during the project, including a new species for the refuge: Blarina hylophaga Elliot (Elliot’s Short-tailed Shrew). The project so far has added 6 species across all taxa to the Table 4. Herpetofaunal species list for Tishomingo National Wildlife Refuge. List includes all species that are known to or may occur in the area. Columns include species observed during this study, species that have been found on Tishomingo NWR prior to starting this survey or from opportunistic encounters during/since the survey started, and species with historic records from Johnston County, OK. This Tish. Johnston Scientific name Common name study NWR County Salamanders Ambystoma texanum (Matthes) Small-mouthed Salamander Ambystoma tigrinum (Green) Eastern Tiger Salamander Notophthalmus viridescens Eastern Newt x Frogs and toads Acris blanchardii Blanchard’s Cricket Frog x x x Anaxyrus americanus American Toad x x x Anaxyrus cognatus (Say in James) Great Plains Toad Anaxyrus woodhousii Woodhouse’s Toad x x x Gastrophryne carolinensis Eastern Narrow-mouthed Toad x Gastrophryne olivacea Western Narrow-mouthed Toad x x x Hyla chrysoscelis/ versicolor Gray Treefrog Complex x x x Hyla cinerea Green Treefrog x Lithobates blairi (Mecham, Littlejohn, Plains Leopard Frog Oldham, Brown, and Brown) Lithobates catesbeianus American Bullfrog x x x Lithobates palustris Pickerel Frog x Lithobates sphenocephalus Southern Leopard Frog x x x Pseudacris clarkii (Baird) Spotted Chorus Frog Pseudacris fouquettei Cajun Chorus Frog x x x Pseudacris streckeri Wright and Wright Strecker’s Chorus Frog x x Scaphiopus hurterii Hurter’s Spadefoot x x Lizards Aspidoscelis gularis (Baird and Girard) Common Spotted Whiptail x Aspidoscelis sexlineata Six-lined Racerunner x x x Crotaphytus collaris (Say in James) Eastern Collared Lizard x Ophisaurus attenuatus Cope Slender Glass Lizard x Plestiodon anthracinus (Baird) Coal Skink Plestiodon fasciatus American Five-lined Skink x x x Plestiodon laticeps Broad-headed Skink x x x Plestiodon obsoletus Baird and Girard Great Plains Skink x Plestiodon septentrionalis Baird Prairie Skink x Southeastern Naturalist J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 192 Table 4, continued. This Tish. Johnston Scientific name Common name study NWR County Sceloporus consobrinus Prairie lizard x x x Scincella lateralis Little Brown Skink x x x Snakes Agkistrodon contortrix Copperhead x x x Agkistrodon piscivorus Cottonmouth x x x Cemophora coccinea (Blumenbach) Scarletsnake x Coluber constrictor Racer x x x Coluber flagellum (Shaw) Coachwhip x Crotalus horridus Timber Rattlesnake x x x Diadophis punctatus Ring-necked Snake x x x Haldea striatula Rough Earthsnake x x x Heterodon platirhinos Latreille Eastern Hog-nosed Snake x x Lampropeltis calligaster Yellow-bellied Kingsnake x x x Lampropeltis holbrooki Stejneger Speckled Kingsnake x x Lampropeltis triangulum (Lacapede) Eastern Milksnake Nerodia erythrogastor (Forster) Plain-bellied Watersnake x x Nerodia rhombifer (Hallowell) Diamond-backed Watersnake x x Nerodia sipedon (L.) Common Watersnake x x Opheodrys aestivus (L.) Rough Greensnake x x Pantherophis emoryi Great Plains Ratsnake x x x Pantherophis obsoletus Western Ratsnake x x x Regina grahamii Baird and Girard Graham’s Crayfish Snake Rena dulcis Baird and Girard Texas Threadsnake Sistrurus miliarius Pygmy Rattlesnake x x x Sonora semiannulata Baird and Girard Western Groundsnake x Storeria dekayi Dekay’s Brownsnake x x x Tantilla gracilis Baird and Girard Flat-headed Snake x Thamnophis proximus Western Ribbonsnake x x x Thamnophis sirtalis (L.) Common Gartersnake x Tropidoclonion lineatum (Hallowell) Lined Snake Virginia valeriae Smooth Earthsnake x x Turtles Apalone mutica (LeSueur,) Smooth Softshell x x Apalone spinifera (LeSueur) Spiny Softshell x x Chelydra serpentina (L.) Snapping Turtle x x Deirochelys reticularia (Latreille, in Chicken Turtle Sonnini and Latreille) Graptemys ouachitensis Cagle Ouachita Map Turtle x x Kinosternon subrubrum (Lacepede) Eastern Mud Turtle x x Macrochelys temminckii Alligator Snapping Turtle x x Pseudemys concinna (LeConte) River Cooter x x Sternotherus carinatus (Gray) Razor-backed Musk Turtle x x Sternotherus odoratus (Latreille) Eastern Musk Turtle x Trachemys scripta Pond Slider x x x Terrapene carolina Eastern Box Turtle x x x Terrapene ornata (Agassiz) Ornate Box Turtle x x Crocodilians Alligator mississippiensis Cuvier American Alligator x x Southeastern Naturalist 193 J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 known-species list. An updated herpetofaunal species list of 73 species has been created for the refuge showing the recently found new species (Table 4). This new herpetofaunal baseline data will allow managers to monitor the effects of current management actions and take the herpetofauna community into consideration when deciding on new management actions. Although Tishomingo NWR was originally created with the goal of providing habitat for waterfowl and migratory songbirds, its varied habitats also support an appreciable richness of reptiles and amphibians. Perhaps other refuges in the NWR system have similar but unrecognized value in protecting multiple taxonomic groups. Future studies will hopefully document such patterns and thereby demonstrate added benefits of the NWR system. Acknowledgments We would like to thank Tishomingo National Wildlife Refuge for allowing access and for funding the project. We would also like to thank multiple interns and volunteers for helping conduct the surveys, including D. and A. Roach, T. Catrett, D. Smith, K. Hurst, B. SoRelle, K. Manktelow, D. Behrens, and C. Eichhorn. Literature Cited Alford, R.A., and S.J. Richards. 1999. Global amphibian declines: A problem in applied ecology. Annual Review of Ecology and Systematics 30:133–165. 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. Benjamini, Y., and D. Yekutieli. 2001. The control of false discovery rate under dependency. The Annals of Statistics 29:1165–1188. Carpenter, C.C. 1958. The amphibians and reptiles of the University of Oklahoma Biological Station area in south central Oklahoma. Proceedings of the Oklahoma Academy of Science 36:39–46. Ceballos, G., P.R. Ehrlich, D.B. Anthony, G. Andres, R.M. Pringle, and T.M Palmer. 2015. Accelerated modern human-induced species losses: Entering the sixth mass extinction. Science Advances 1:1–5. Collins, J.P., and A. Storfer. 2003. Global amphibian declines: Sorting the hypothesis. Diversity and Distribution 9:89–98. Conant, R., and J.T. Collins. 1998. A Field Guide to Reptiles and Amphibians: Eastern and Central North America. Vol. 12. Houghton Mifflin, Boston, MA. Corn, P.S. 1994. Straight-line drift fences and pitfall traps. Pp. 109–117, In W.R. Heyer, M.A. Donnelly, R.W. McDiarmid, and L.C. Hayek (Eds.). Measuring and Monitoring Biological Diversity: Standard Methods for Amphibians. Smithsonian Institution Press, Washington, DC. 384 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. Enge, K.M., and Marion, W.R. 1986. Effects of clearcutting and site preparation on herpetofauna of a North Florida flatwoods. Forest Ecology and Manag ement 14:177–192. Southeastern Naturalist J.A. Muller, J.A. Veech, and J. Roach 2017 Vol. 16, No. 2 194 Fitch, H.S. 1992: Methods of sampling snake populations and their relative success. Herpetological Review 23:17–19. Geluso, K., and C.H. Jones. 2004. Westward expansion of the Eastern Fox Squirrel (Sciurus Niger) in northeastern New Mexico and southeastern Colorado. Southwestern Naturalist 49:111–116. Gibbons, J.W., and R.D. Semlitsch. 1982. Terrestrial drift fences with pitfall traps: An effective technique for quantitative sampling of animal populations. Brimleyana 7:1–16. Hanlin, H.G., F.D. Martin, L.D. Wike, and S.H. Bennett. 2000. Terrestrial activity, abundance, and species richness of amphibians in managed forests in South Carolina. American Midland Naturalist 143:70–83. Hayes, T.B., P. Case, S. Chui, D. Chung, C. Haeffele, K. Haston, M. Lee, V.P. Mai, Y. Marjuoa, J. Parker, and M. Tsui. 2006. Pesticide mixtures, endocrine disruption, and amphibian declines: Are we underestimating the impact? Environmental Health Perspectives 114:40–50. International Union for the Conservation of Nature (IUCN). 2008. Red list of threatened species. Available online at http://www.iucnredlist.org/initiatives/amphibians/analysis/ red-list-status. Accessed 24 March 2015. Krebs, C.J. 1999. Ecological Methodology, 2nd Edition. Addison-Wesley, Menlo Park, CA. 654 pp. Petranka, J.W., M.E. Eldridge, and K.E. Haley. 1993. Effects of timber harvesting on southern Appalachian salamanders. Conservation Biology 7:363–370. 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. Renken, R.B., W.K. Gram, D.K. Fantz, S.C. Richter, T.J. Miller, K.B. Ricke, B.Russell, and X. Wang. 2004. Effects of forest management on amphibians and reptiles in Missouri Ozark forests. Conservation Biology 18:174–188. Sievert, G., and L. Sievert. 2005. Field Guide to Oklahoma's Amphibians and Reptiles. Oklahoma Department. of Wildlife Conservation, Oklahoma City, OK. 211 pp. Sokal, R.R., and F.J. Rohlf. 1995. Biometry: The Principles and Practice of Statistics in Biological Research, 3rd Edition. W.H. Freeman and Co., New York, NY. 880 pp. Todd, B.D., C.T. Winne, J.D. Willson, and J.W. Gibbons. 2007. Getting the drift: Examining the effects of timing, trap type, and taxon on herpetofaunal drift fence surveys. American Midland Naturalist 158:292–305. US Fish and Wildlife Service (USFWS). 2010. Tishomingo National Wildlife Refuge: Comprehensive conservation plan. Available online at http://www.fws.gov/southwest/ refuges/Plan/docs/Oklahoma/Tishomingo_CCP/index.html. Accessed 2 March 2015. 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. White, J.A., P.R. Moosman, C.H. Kilgore, T.L. Best, and P.D. Sudman. 2006. First record of the Eastern Pipistrelle (Pipistrellus subflavus) from Southern New Mexico. Southwestern Naturalist 51:420–422. Wilson, E.O. 1988. The current state of biological diversity. Pp. 3–18, In E.O. Wilson (Ed.). Biodiversity. National Academy Press, Washington, DC. 521 pp. Zar, J.H. 1999. Biostatistical Analysis, 4th Edition. Prentice Hall, Upper Saddle River, NJ. 663 pp.