Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 513 2018 NORTHEASTERN NATURALIST 25(4):513–531 Observations of Wood Turtle Activity, Diet, Movements, and Morphometrics in the Central Appalachians Kathryn R.P. McCoard1, Noah S. McCoard1,2, and James T. Anderson1,* Abstract - Glyptemys insculpta (Wood Turtle) populations are declining as habitat is developed and individuals are collected for the pet trade. Information about the natural history and life history of the species can help inform conservation strategies, especially for populations about which little information is currently available. Because basic ecological information is lacking for populations in the Central Appalachians, we studied the natural history of Wood Turtles in West Virginia from 2009 to 2011. Wood Turtle males were larger than females in carapace length and width, bridge height, and mass. Turtles were primarily terrestrial in spring and summer and aquatic in autumn and winter. Aquatic mating was most prevalent in autumn. Nesting attempts, which were mostly in sandy substrates, were made in spring during early morning and evening. Slugs made up the majority (67%) of the turtles’ diet, although other invertebrates, vertebrate remains, berries, and green leaves were also consumed. In autumn, turtles entered the river for brumation. This study adds to our understanding of the natural history of Wood Turtles near the southern extent of their range. Introduction Glyptemys insculpta (LeConte) (Wood Turtle) is an endangered species due primarily to habitat loss, habitat fragmentation, inadequate recruitment, and collection for the pet trade (Daigle and Jutras 2005, IUCN 2011, Saumure et al. 2007). The range of the Wood Turtle extends from eastern West Virginia and northern Virginia north through the Atlantic states to Nova Scotia, Canada, and west to eastern Minnesota (Ernst and Lovich 2009). In West Virginia, Wood Turtles are listed as a vulnerable species in need of conservation (WVDNR 2015). They are under review for listing under the US Endangered Species Act (USFWS 2017). Moreover, international trade of Wood Turtles is monitored through the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES 2008). Although there have been several studies of Wood Turtles in West Virginia, gaps in our knowledge remain (Breisch 2006; Curtis and Vila 2015; McCoard et al. 2016a, b; Niederberger 1993; Niederberger and Seidel 1999). In particular, more information is needed on diet, activity cycles, and movements of Wood Turtles near the southern extent of their range. As part of a radio telemetry (McCoard et al. 2016a) and population (McCoard et al. 2016b) study on Wood Turtles in the eastern panhandle of West Virginia, we had the opportunity to 1West Virginia University, School of Natural Resources, PO Box 6125, Morgantown, WV 26506. 2Current address - Georgia Department of Natural Resources, Wildlife Resources Division, 703 E. Ward Street, Douglas, GA 31533. *Corresponding author - jim.anderson@mail.wvu.edu. Manuscript Editor: Todd Rimkus Northeastern Naturalist 514 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 Vol. 25, No. 4 observe behavioral and natural history attributes of this species. Our objective was to record natural history observations of Wood Turtles in West Virginia, including morphometrics, activity cycles, and diet. Field-Site Description We recorded natural history observations of Wood Turtles along a 14-km reach of a stream in the eastern panhandle of West Virginia. Maximum water depths reached 2.5 m and the stream varied from 10 to 30 m wide. A diversity of native warmwater fish species occupy the stream. Portions of the stream have highly eroded streambanks reaching 2–3 m in height; however, other areas have intact banks with little erosion. Primary land use was dominated by agriculture, primarily hay fields, cornfields, and cattle pastures, but also included eastern deciduous forests. A baseline assessment of the watershed was conducted by Constantz et al. (1995). The eastern panhandle of West Virginia is within the Ridge and Valley Province and receives 76 cm of precipitation annually (Kozar and Mathes 2001). Average annual frost-free period is at least 180 days (Curry 1978). Methods Observations We surveyed on foot or by canoe 5 locations located along the 14-km stretch. Our surveys, conducted from May 2009 to August 2011, varied from 600 to 1100 m of river length. We surveyed perpendicularly from either side of the river’s edge to 150 m (a distance greater than 95% of freshwater turtle migrations measured by Bodie [2001] and 90% of radio-telemetered Wood Turtle movements observed in Québec, Canada by Arvisais et al. [2002]). The sites were 300 to 1000 m apart from their neighboring sites. Upon encountering Wood Turtles, we attempted to remain inconspicuous and document their activities and behaviors. Once they responded to our presence, or were otherwise disturbed, we would capture turtles by hand or dipnet. We captured turtles under permits from the West Virginia Division of Natural Resources and with approval from the West Virginia University Animal Care and Use Committee (protocol # 09-0408). We uniquely marked adult turtles by filing notches into the marginal scutes of the carapace with a triangular file (Cagle 1939). Juveniles less than 2 yrs. old were marked with white enamel paint to avoid physical alteration (McCoard et al. 2016a). On a subset of Wood Turtles (n = 31; 15 males, 10 females, and 6 juveniles), we affixed radio transmitters (ATS R1860, 15 g, < 10% of turtle mass; Advanced Telemetry Systems, Isanti, MN) with epoxy to the back right edge of the carapace. We tracked them between the hours of 0800 and 1930 once to twice per week during March to October, and once per month from November to February each year (McCoard et al. 2016a). Telemetry facilitated additional opportunistic Wood Turtle observations. Upon initial capture of each turtle, we measured to the nearest millimeter minimum straight-line carapace length (CL), maximum carapace width (CW), minimum straight-line plastron length (PL), maximum plastron width (PW), maximum bridge height (BH) and width (BW), and maximum depth from carapace top to plastron Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 515 bottom (D) using 200-mm (± 0.2 mm) MitutoyoTM Dial Calipers (Mitutoyo America Corp., Aurora, IL). We sexed individuals that were ≥160 mm CL based on secondary sexual characteristics (Harding and Bloomer 1979). We measured mass using 1000-g (± 10 g) or 2500-g (± 20 g) Pesola® spring scales (Pesola Präzisionswaagen AG, Schindellegi, Switzerland). To estimate turtle’s age, we counted growth rings on the carapace from multiple scutes and recorded the most consistent number of rings obtained (Harding and Bloomer 1979), although accuracy of the method is unreliable for some species (Brooks et al. 1997, Litzgus and Brooks 1998). The rings begin to fade after ~20 years, and in such cases age was recorded as >20 years. For every capture, we recorded date, time, global positioning system (GPS, Garmin eTrex Legend, ± 3–6 m; Garmin Ltd., Olathe, KS) location, weather conditions, activity when observed or captured, identification marks, and perpendicular distance from the river. We released all Wood Turtles at their points of capture after they were processed. Microclimate data We collected microclimatic data to attempt to determine conditions that were preferred by Wood Turtles for their various activities in West Virginia (Ernst 1986, Reagan 1974). If the captured turtles were terrestrial, we collected the data directly under them to gain close approximates of the environment being used, including a single reading of soil temperature (ST; ± 1% of scale; soil thermometer; Forestry-Suppliers, Jackson, MS), soil pH (SpH; ± 0.01; Oakton® Double Junction Waterproof pH Tester 30; Oakton Instruments, Vernon Hills, IL), and soil moisture (SM; 1 = dry, 10 = saturated; 23-cm Lincoln Soil Moisture Meter; Lincoln Irrigation Co., Lincoln, NE). If the turtles were aquatic, we measured water temperature (WT; °C; 15-cm Enviro-Safe® Armor Case Pocket Thermometer; Bel-Art H-B Instrument, Wayne, NJ). We also recorded air temperature (AT; ± 1 °C) either next to terrestrial turtles or above the water’s surface directly over aquatic turtles. Statistical analyses We performed all statistical analyses in R 2.10.1 with α = 0.05 (R Development Core Team). We used Welch 2-sample t-tests to compare morphometric data among males and females in the overall population, accounting for unequal variances and unequal sample sizes. We used paired t-tests to compare morphometric data among males and females observed mating. Statistical analyses were not conducted on juvenile measurements. To determine if the number of terrestrial and aquatic mating observations differed between autumn and spring, we used a Pearson chi-square (χ²) test, which assumes independent observations, replicated data, and sufficient sample sizes (Kutner et al. 2005). To determine if the number of males, females, and juveniles differed in being terrestrial or aquatic based on season, we used a contingency table with a χ² test. If a significant marginal (over all sexes) association was found, we used a Fisher’s exact test, which tests the independence of rows and columns of the contingency table, to determine if a significant conditional (within sex) as sociation existed. Northeastern Naturalist 516 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 Vol. 25, No. 4 Results Morphometrics We captured 284 unique Wood Turtles (males = 137, females = 88, and juveniles = 59). Total captures numbered 1443 (1159 recaptures, 80.3%). Mean PL, PW, BW, and D did not differ between adult males and females (P > 0.05); however, males were larger in CL, CW, BH, and mass than females (Tables 1, 2). Mating males were larger than mating females in CL, BH, and mass (P < 0.05). Five turtles (2 males, 2 females, and 1 juvenile) were found dead by unknown causes during the study. Nineteen turtles had punctures and deep cuts on their carapaces and an additional 15 turtles were missing appendages or an eye. Activity cycles Aquatic and terrestrial habitat use. Season influenced whether Wood Turtles (males, females, and juveniles, collectively) were terrestrial or aquatic (χ3² = 245.54, P < 0.001; Table 3), and locations appeared to differ depending on environmental conditions (Table 4). The overall trend indicated high terrestrial activity in spring and summer, slightly higher aquatic activity in autumn, and an obligatory aquatic stage in winter. Males were primarily terrestrial in spring and summer and aquatic in autumn and winter (P < 0.001). Females were almost equally aquatic and terrestrial in spring and autumn, but predominantly terrestrial in summer and aquatic in winter (P < 0.001). Juveniles followed the same trend as males, with more equal proportions in autumn (P < 0.001). Emergence. We observed the first turtle emerging from brumation, a period of inactivity due to cold temperatures, in 2010 on 12 March. Temperatures were as follows: ST = 10 °C; AT = 12 °C; and WT = 8 °C. In 2011, the first turtle was seen out of brumation on 19 March when ST = 13 °C, AT = 26.4 °C, and WT = 11 °C. Both turtles were male and within 1 m of the river’s edge, basking. From these dates, turtles became active, primarily remaining within or near the river as spring mating began. Mating. Mating occurred in the spring from late March to early June (n = 20) when mean temperatures were as follows: ST = 11.5 °C (SE = 0.84; min–max = 7–17 °C); AT = 13.2 °C (SE = 1.36; min–max = 7.1–27.3 °C); and WT = 11 °C (SE = 0.57; min–max = 8–16 °C). Autumn mating occurred from late August to early November (n = 36) when mean temperatures were as follows: ST = 14.1 °C (SE = 0.73; min–max = 8–21 °C); AT = 18.8 °C (SE = 0.87; min–max = 8.8–27.9 °C); and WT = 15.1°C (SE = 0.85; min–max = 8–24 °C); slightly warmer than the average spring temperatures. Twenty-eight mating pairs (n = 45 individuals) were observed; 8 (17.8%; 5 females and 3 males) of the individuals were found mating more than once (with different partners except in 1 case) between autumn 2009 and spring 2011. Autumn mating accounted for 64.3% of all mating observations, and 75% of all mating observations occurred after 1300 hrs (earliest–latest = 0920–1814 hrs). Of the mating pairs, 10 (35.7%) were terrestrial at an average distance of 13.5 m (SE = 3.06; min–max = 0–30 m) from the river’s edge; 90% of those mating attempts were in autumn. When mating was aquatic, 18 (64.3%) turtle pairs were an Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 517 Table 2. Means, standard errors (SE), minimum, and maximum of each morphometric variable recorded for all Wood Turtles observed mating (n = 25 males, n = 18 females) along a 14-km stream reach in the eastern panhandle of West Virginia, from spring 2009 through summer 2011. All measurements are in mm, except mass (g). Within a row, means followed by the same letter did not differ between males and females observed mating. Mating males Mating females Variable Mean SE Min Max Mean SE Min Max t df P Carapace width 146.46a 1.38 134.38 165.10 142.84a 1.58 128.00 155.55 1.72 37.48 0.094 Plastron length 180.77a 3.58 113.79 206.00 179.25a 4.31 115.34 204.27 0.27 36.28 0.789 Plastron width 113.12a 1.97 74.50 124.00 119.72a 3.27 108.04 172.42 1.73 28.86 0.095 Bridge width 69.51a 1.24 56.80 84.55 68.25a 1.25 61.92 81.24 0.71 39.69 0.482 Bridge height 24.24a 0.38 20.47 28.00 22.33b 0.36 20.00 25.58 3.66 40.41 less than 0.001 Depth 69.05a 0.83 58.00 78.00 68.31a 0.75 62.53 72.00 0.66 40.80 0.516 Mass 1154.00a 24.75 910.00 1400.00 1043.00b 27.72 920.00 1300.00 2.99 37.87 0.005 Table 1. Means, standard errors (SE), minimum, and maximum of each morphometric variable recorded for all unique Wood Turtles (n = 284 total, n = 137 males, n = 88 females, n = 59 juveniles) captured along a 14-km stream reach in the eastern panhandle of West Virginia, from spring 2009 through summer 2011. All measurements are in mm, except mass (g). Within a row, means followed by the same letter did not differ between males and females in the overall population. Statistical analyses were not done on juvenile measurements. Males Females Juveniles Variable Mean SE Min Max Mean SE Min Max Mean SE Min Max t df P Carapace length 194.6a 1.28 160.0 232.2 180.7b 1.18 167.6 206.9 102.1b 4.27 36.6 151.0 8.00 218.65 less than 0.001 Carapace width 141.7a 1.01 111.4 183.6 136.9b 1.25 103.0 191.1 82.8b 2.79 36.0 116.2 2.94 186.13 0.004 Plastron length 178.2a 1.10 109.0 206.0 175.5a 1.82 108.3 204.3 99.2b 4.45 34.2 152.9 1.30 148.91 0.197 Plastron width 114.2a 0.95 95.0 197.4 116.9a 1.57 90.8 192.3 65.9b 2.48 26.3 93.9 1.47 150.06 0.144 Bridge width 68.8a 0.58 50.7 95.0 68.1a 0.65 53.0 87.8 34.9b 1.65 10.1 54.9 0.84 198.48 0.405 Bridge height 23.6a 0.23 13.5 36.5 21.4b 0.21 15.5 26.6 11.6b 0.46 5.0 20.0 7.15 217.54 less than 0.001 Depth 67.8a 0.50 46.0 79.6 67.7a 0.52 52.1 79.5 37.1b 1.51 9.4 56.1 0.18 208.41 0.855 Mass 1116.0a 15.82 520.0 1430.0 981.1b 18.15 430.0 1380.0 198.0b 19.37 7.0 500.0 5.65 197.51 less than 0.001 Northeastern Naturalist 518 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 Vol. 25, No. 4 average of 2.64 m (SE = 0.71; min–max = 0–10 m) from the shore and in water that was 0.5 m (SE = 0.066; min–max = 0.25–1 m) deep; 50% of those observations occurred in autumn. All aquatic pairs were seen in quiet pools or adjacent to the river’s main current. The number of terrestrial and aquatic mating attempts was not different with respect to season (i.e., spring and autumn; χ1² = 0.034, P = 0.853). When the turtles were observed attempting mating, the male would wrap his front and back claws under the lip of the female’s carapace and hold on. When the female was unresponsive, the male would push up from the female and slam his plastron down on her carapace. The male would extend his neck and peer down at the female, biting her if she tried to extend her neck or flee. We observed no actual copulation, and the majority of males released the female upon noticing us. Occasionally, the male would pursue the female soon after our disruption. Nesting and nest emergence. We made 3 observations of females digging multiple nests, but not to completion. The nest holes were made in late May to early June in the early morning and early evening. Throughout this period, many small holes were observed along the stream banks that appeared to be nest attempts; however, we did not observe turtles creating all of the holes. All uncompleted nests typically terminated in substrate that was too rocky. The nest sites were sandy, intermixed with pebbles, with little to no vegetation; they were typically 0.3–1 m above water level and less than 15 m from the river ’s edge. We made detailed observations of nesting of one female. She was in a trancelike state as she laid her eggs. She used one back foot to brace herself and the other to catch the eggs and maneuver them into the nest chamber. After laying the final egg, as she filled in the nest, alternating her hind feet as she scooped the sand, any movements we made were instantly noticed by her, causing her to pause. Once the nest was covered, she moved to the water. This nesting event occurred on 24 May 2010 at 0930 hrs during a light rain. The female’s measurements were: CL = 181.3 mm, CW = 126.12 mm, PL = 179.31 mm, PW = 121.6 mm, BW = 63.4 mm, BH = 22.1 mm, D = 67.45 mm, and post-laying mass = 980 g. She was >20 years old. She nested 6 m from the side of the river, at the edge of vegetation. The nest was 140 mm deep, 120 mm wide, and 65 mm deep to the topmost egg. The female laid 11 eggs with an average length of 33 mm (SE = 0.11; min–max = 32.2–33.5 mm), width of 24 mm (SE = 0.05; min–max = 23.7– Table 3. Proportions of observations in each season for the overall Wood Turtle (n = 1443 captures) population and of males (n = 751), females (n = 524), and juveniles (n = 168) based on terrestrial (T) or aquatic (A) occurrence along a 14-km stream reach in the eastern panhandle of West Virginia from spring 2009 through summer 2011. Population Male Female Juvenile Season A T A T A T A T Spring 0.36 0.64 0.25 0.75 0.46 0.54 0.33 0.67 Summer 0.23 0.77 0.31 0.69 0.14 0.86 0.14 0.86 Autumn 0.59 0.41 0.63 0.37 0.52 0.48 0.58 0.42 Winter 0.99 0.01 0.98 0.02 1.00 0.00 1.00 0.00 Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 519 Table 4. Environmental variables (mean, standard error, and min - max) related to Wood Turtles (n = 1419 captures) being aquatic (A) or terrestrial (T) depending on season along a 14-km stream reach in the eastern panhandle of West Virginia from spring 2009 to summer 2011. Headings are as follows: soil temperature (ST), air temperature (AT), water temperature (WT), and soil moisture (SM; scale of 1 [dry] to 10 [saturated]). ST (°C) AT (°C) WT (°C) SM Season Location Mean SE Min Max Mean SE Min Max Mean SE Min Max Mean SE Min Max Spring A 12.3 5.0 5.0 27.0 16.2 8.3 5.1 35.5 12.0 4.4 7.0 25.5 6.5 2.0 1.0 10.0 T 19.7 4.3 10.0 33.0 26.7 6.5 12.2 40.6 18.2 3.7 11.0 27.0 6.0 2.6 1.0 10.0 Summer A 23.4 3.1 17.0 34.0 29.2 4.1 20.0 42.0 23.6 2.6 19.0 29.0 4.3 2.5 1.0 10.0 T 22.6 2.8 16.0 31.0 29.8 11.0 18.5 46.0 24.2 2.7 17.0 34.0 3.9 2.8 1.0 10.0 Autumn A 10.6 6.3 -2.0 23.0 14.6 5.7 1.6 27.0 10.3 3.7 1.0 19.0 4.9 2.6 1.0 10.0 T 13.9 3.7 7.0 25.0 18.0 5.6 5.7 29.5 13.6 3.5 7.0 18.5 5.3 2.6 1.0 10.0 Winter A 4.8 4.3 0.0 13.5 9.3 6.9 -3.0 34.4 4.6 3.8 0.0 11.0 3.2 3.1 1.0 10.0 T 5.0 3.5 0.0 10.0 4.7 5.2 -2.7 12.0 4.0 2.8 0.0 8.0 4.5 2.5 1.0 8.0 Northeastern Naturalist 520 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 Vol. 25, No. 4 24.2 mm), and mass of 15 g (SE = 0.04; min–max = 14.8–15.2 g). Soil temperature was 20 °C, AT was 19.1 °C, and WT was 18 °C. The soil had a low moisture rating (SM = 2) and the pH was 7.3. Canopy cover was 0%. We observed 3 hatchlings emerge from the nest on 25 July 2010, almost exactly 2 months later, at ~1730 hrs. Their means were: CL = 37.98 mm (SE = 1.04; min– max = 36.56–40.02 mm), CW = 38.09 mm (SE = 1.08; min–max = 36.02–38.58 mm), PL = 34.33 mm (SE = 0.11; min–max = 34.17–34.53 mm), PW = 27.39 mm (SE = 0.69; min–max = 26.28–28.66 mm), BW = 10.38 mm (SE = 0.16; min–max = 10.07–10.56 mm), BH = 5.32 mm (SE = 0.23; min–max = 5.04–5.78 mm), D = 15.7 mm (SE = 0.19; min–max = 15.46–16.08 mm), and mass = 9.33 g (SE = 0.33; min– max = 9–10 g). Soil temperature was 23 °C, AT was 22.7 °C, and WT was 23 °C. Soil moisture, SpH, and canopy cover remained the same. Soon after emergence, the hatchlings began consuming their eggshells. Active season. When turtles were aquatic, we saw them walking along the riverbed, exploring log jams and root masses, and poking their heads up for a breath. Although swimming was observed, it was less common than walking along the bottom. Within the river on warm days, the turtles often took refuge in root masses protruding from the bank, thick organic mud on the streambed, fallen logs, leaf litter, or undercut banks. During these days, AT varied from 27.3 to 42 °C (mean = 31.4 °C, SE = 0.34), ST was 19–34 °C (mean = 23.8 °C, SE = 0.32), and WT was 19–29°C (mean = 24.1 °C, SE = 0.26). When the turtles were terrestrial (AT = 12.2–46 °C; ST = 10–33° C; WT = 11–34 °C), we often found them walking along Odocoileus virginianus (Zimmermann; White-tailed Deer) trails bordering the bank, through woods, and up mountainsides. On 20 July 2009, an adult male Wood Turtle was encountered walking on a deer trail running parallel to the river. When he came across an intersecting deer trail running perpendicular to the river, he turned left, away from the river, and began following the intersecting trail. Clear paths were not always chosen for travelling, however. Thick grasses and Verbesina alternifolia L. (Wingstem) reaching 1–2 m tall did not deter the turtles from forging along the ground. When turtles were not found walking about during the day, we often found them sitting still, legs tucked in and head out, in sparse vegetation. We typically found turtles estivating, inactive and burrowed during warmer temperatures, in the spring (26.5% of all terrestrial observations) and fall (27.5%). Nearly half (45%) of all terrestrial observations in the summer were of turtles estivating, when AT varied from 20.9 to 40 °C (mean = 29.6 °C, SE = 0.44), ST was 17 to 29 °C (mean = 22.4 °C, SE = 0.31), and WT was 19–33 °C (mean = 23.9 °C, SE = 0.33) Locations included matted grasses (dead and alive), thick herbaceous cover, thick shrubs (especially Rosa multiflora [Thunb.] [Multiflora Rose]), undercuts along banks, and leaf litter piled against woody debris. Occasionally, we found individuals below the surface in deep hoof depressions created by Bos taurus (L.) (Domesticated Cattle) in a saturated area of pasture. Basking. We observed basking turtles during all seasons except winter. Of the basking events we observed, 61% occurred in the spring. Although the turtles were Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 521 seen basking from 0821 to 1657 hrs, 58% of our observations took place before noon. Basking sites included streambanks, deer trails on mountainsides, clearings in vegetation, rocky beaches, and propped up against matted grasses and woody debris to achieve an approximate 45° angle towards the sun. Their heads and appendages were usually extended. During basking, ST, AT, and WT varied from 8 to 33 °C (mean = 20.4 °C, SE = 0.55), 8 to 40 °C (mean = 27.6 °C, SE = 0.72), and 7 to 27° C (mean = 18.3 °C, SE = 0.57), respectively. Canopy cover was 0–100% (mean = 36.5%, SE = 0.03). Diet. During the active season, we made dietary observations on 70 occasions. Initial observations occurred in April, at which time turtles ate slugs and unidentified green leaves. A single male was observed repeatedly slamming his body down against the ground, a behavior identified as stomping for worms (Kaufman 1986). In May, common foods consumed included Impatiens spp. (jewelweed), slugs, and worms. The largest variety of items eaten and the greatest number of eating observations (47%) occurred in June and included jewelweed, Arisaema dracontium ([L.] Schott) (Green Dragon), Phyllophaga spp. (June bugs), slugs, worms, and unidentifiable small mammal remains. On an overcast day, we observed a female worm-stomping. She was standing in a clearing on muddy ground, not long after rain had ended, about 10 m from the river’s edge. Her front right leg was wiggled back and forth as her foot was pressed against the ground, followed by her front left leg and foot. Suddenly she began rapidly raising the front end of her body up, slamming into the ground repeatedly. While observed, she did not catch any worms, but she quickly became conscious of being watched and ended her routine. In July, only slugs were found being eaten. Slugs, Phytolacca americana L. (Pokeweed) berries, Elaeagnus umbellata (Thunb.) (Autumn Olive) berries, Taraxacum officinale (F.H. Wigg.) (Dandelion) leaves, and unidentifiable songbird remains made up the prey for August. We made final diet observations in September. During this month, Pokeweed berries, green leaves, Viola spp. (violets), and Prunus serotina (Ehrh.) (Wild Black Cherry) fruits were consumed. Overall, slugs were the most frequently consumed (67%) food. All other items made up less than 1% of the diet observations. Dominance and aggression. We observed dominance displays on a couple of occasions, primarily within the river. On 3 October 2009, we saw a male chasing another male during the fall mating season. Two males were observed nudging each other with their heads on 7 November 2009. A male was found mounted on another male on 1 October 2010. On 19 November 2010, around the time that the turtles were entering into hibernation, a non-radio-tagged male approached a radio-tagged male in the latter’s home range (McCoard et al. 2016a), their necks outstretched. The resident male attempted a bite, causing the other to flee. A single terrestrial dominance display occurred on 14 June 2010; a radio-tagged male was found in his home range (McCoard et al. 2016a) next to a new, unmarked male. When the new male tried to walk away, the resident male bit his front leg. The new male paused, tucking into his shell as the resident male stood with his neck outstretched above him. As the new male again tried to leave, the resident male pursued him, biting. Northeastern Naturalist 522 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 Vol. 25, No. 4 The sequence of aggression appeared to follow this order: the dominant individual bit at the subordinate’s eyes first, then the legs as the latter’s head was pulled into his shell; the resident individual then asserted his dominance by fully extending his neck vertically; if the new male raised his head in the presence of the resident individual, the resident opened his mouth for a few seconds, then started trying to bite the new male again. This continued until the new male, who we concluded to be the subordinate individual, was able to successfully flee. Brumation. Turtles began spending the majority of their time in the river by late October, continuing to move about, and thus not yet brumating. However, by early November, the majority of turtles were in brumation (n = 163 observations). At this time, ST was 7–9 °C, AT was 12.2–14.5 °C, and WT was 7 °C. During brumation, mean environmental temperatures were as follows: ST = 1.65 °C (SE = 0.26; min–max = -2–9 °C), AT = 6.42 °C (SE = 0.46; min–max = -3–15.1 °C), and WT = 3.77 °C (SE = 0.34; min–max = 0–8 °C). Typical brumation sites were long, quiet pools that reached a depth of ~1– 2.5 m and width of about ~20– 30 m. Turtles in hibernations were on average 0.78 m (SE = 0.05, min–max = 0.25–2.50 m) below the surface and 3.39 m (SE = 0.47, min–max = 0.25–20.00 m) from shore. Turtles displayed communal brumation in the same pools, but were usually spaced >1 m apart. Turtles were either covered in soft organic substrate, sand, or leaf litter; in the root mass of a fallen tree or one protruding from the bank; or sitting exposed on the riverbed. Twenty-six of the turtles were found under 2.5–10.0 cm thickness of ice that spanned the river, with the nearest unfrozen surface water up to 30 m away. Two Wood Turtles were found covered in algae so thick that it was hard to distinguish them from the surrounding rocks. We saw little movement by brumating turtles except during occasional warm spells when the turtles became sluggishly active. In February 2011, the weather fluctuated between warm and cold periods, and turtles began moving when WT was ~10 °C. Temperature fluctuations continued into early March. One radio-tagged female moved toward shore during an unusually warm spell (AT = 19.4 °C) that lasted about 3 days from late February into early March, after which the air temperature dropped to below freezing (AT < 0.0 °C). We found the female dead in the water, at the shore’s edge, with all of her legs extended. With the return of freezing temperatures, the turtles stopped moving and began brumating again; water temperatures at the time were 7–8 °C. Discussion Morphometrics Our apparently youngest identifiable male had 7 growth rings and a CL of 161.6 mm. Our youngest female had 9 growth rings and a CL of 167.6 mm. The youngest reproductively active male and female were estimated to be 15 and 16 (based on growth rings), respectively. Comparatively, the youngest identifiable males in a Québec agri-forest study were 10 years old with a CL of 176.3 mm (agricultural site) and 11 years old with a CL of 157.6 mm (forested site); the youngest nesting female (forested site) was 15 years old with a CL of 195.4 mm (Saumure and Bider Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 523 1998). In New Jersey, secondary sexual characteristics began to appear when turtles were about 9 years old; maturity was achieved with reproductive activities about 14 years old (Farrell and Graham 1991). In Vermont, a radio-tracked juvenile male, tagged at 2 years old, was observed again at 11 years old with a concave plastron (a secondary sexual characteristic) and a CL of 173 mm; the youngest male found mating was 15 years old (Parren 2013). Male Wood Turtles in our study were longer, thicker, and heavier than the females with whom they mated. Large males may be capable of defending home ranges in prime territory from smaller males, possibly providing them with access to better foraging and more females. Males that are larger than females may be harder for reluctant females to dislodge during mating and, thus, may be more reproductively successful once the female’s resistance breaks down. Male Wood Turtles commonly have longer carapaces than females as observed in New Jersey (Farrell and Graham 1991), Virginia (Akre 2002), and another West Virginia population (Breisch 2006). Although the body sizes of endotherms tend to be larger at higher latitudes (Bergmann’s rule, Northern Hemisphere), ectotherms do not necessarily follow that trend (Angielczyk et al. 2015, Litzgus et al. 2004). The measurements of our Wood Turtles in West Virginia (southern geographic range) fall between the sizes of a New Jersey population (middle geographic range; Farrell and Graham 1991) and a Québec population (northern geographic range; Saumure et al. 2007). Our finding reflects those of previous studies in which turtles in northern (Saumure and Bider 1998) and southern populations were larger, on average, than those in the middle of the range (Greaves and Litzgus 2009, Litzgus et al. 2004, Verdon and Donnelly 2005). Larger sizes in southern turtle populations may result from longer growing seasons, allowing turtles to forage for longer periods of time than their mid-latitude counterparts; larger sizes in northern populations may be a result of greater caloric intake during intensive summer foraging coupled with a longer brumation period of low energy expenditure which results in increased growth. Wood Turtles in our site were regularly found in cornfields, hayfields, and cattle pastures (McCoard et al. 2016a) and traversed their home ranges even as heavy machinery, used on river restoration efforts, passed by them (McCoard 2012). Indeed, a lack of vegetation in harvested hayfields facilitates straight line travel, potentially reducing travel time across exposed fields (Saumure et al. 2010). In Québec, turtles on agricultural sites grew to smaller sizes than their forest counterparts (Saumure and Bider 1998). IThe smaller sizes were thought to be due to mutilation of the turtles by agricultural activities (which also may be the case in our study area): limb loss reduced foraging abilities and shell damage reduced shell growth (Saumure and Bider 1998). Larger turtles also may be more susceptible to mortality in agricultural settings (Tingley et al. 2009). Interestingly, our turtles, living in an agri-forest environment had similar mean sizes and ranges to the Québec agriculture turtles. Similar turtle sizes at our site and the Québec site may be due, in part, to limited food, water, and cover in the narrow riparian zones, which limits growth potential of the turtles. Northeastern Naturalist 524 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 Vol. 25, No. 4 Seasonal activity cycles Aquatic and terrestrial habitat use. During spring, activities in our population were mainly aquatic (mating), but the turtles soon left the river to nest and return to their annual home ranges (McCoard et al. 2016a). Fewer females were terrestrial in the spring than males, possibly related to the females using the river as a refuge between nesting attempts. In summer, individuals were predominantly terrestrial; an observation also noted in Minnesota (Brown et al. 2016) and Pennsylvania (Ernst 1986, Kaufmann 1992a). By autumn, most individuals in our population returned to the river for mating and eventual brumation. In winter, brumation was aquatic as documented by others (Farrell and Graham 1991, Greaves and Litzgus 2007, Harding and Bloomer 1979). Emergence. Wood Turtles in our population emerged from brumation in mid- March, earlier than their northern counterparts, but during similar environmental temperatures. In Ontario, Canada, at the northern extent of the Wood Turtles’ range, the turtles emerged from brumation in mid-April when temperatures were: AT = 13.5 °C and WT = 5.0 °C (Greaves and Litzgus 2007); similar observations were made in Vermont (Parren 2013). Turtles returned to the stream during nights when AT was ≤10 °C and during days when AT was ≤20 °C following emergence in Pennsylvania (Kaufmann 1992a). Mating. Courtship followed soon after emergence (late March to early June) and again in late August to early November. Autumn was the primary mating season for our population and a Québec (Walde et al. 2003) and Vermont population (Parren 2013). Males were more aquatic than females in autumn, which may possibly be a reproductive strategy: the sooner a male arrives, the better chances he may have to mate with a number of females as they trickle in before other males arrive. Of our study’s mating attempts, 35.7% were terrestrial and up to 30 m from the river’s edge, uncommon for Wood Turtles that primarily have aquatic mating (Ernst 1986). In Vermont only 4 of 57 (7.0%) mating attempts occurred on land, with none farther than 18.3 m from the stream (Parren 2013). In Pennsylvania, AT was 11–22.8 °C and ST and WT were 10.0 °C and 20.0 °C, respectively, during mating (Ernst 1986). Mating occurred from mid-April to mid-May and late August to October in Pennsylvania (Kaufmann 1992a), mid-June in Algonquian Park, Canada (Quinn and Tate 1991), May to November in Québec (Walde et al. 2003), and late March to April and October to November in New Jersey (Farrell and Graham 1991). Nesting and nest emergence. Following courtship, nesting was observed during late May to late June in our study, Virginia (Akre 2002), and Vermont (Parren 2013); late May to mid-June in New Jersey (Castellano et al. 2008); and June in Pennsylvania, New Jersey, Michigan, and Québec (Arvisais et al. 2002, Farrell and Graham 1991, Harding and Bloomer 1979, Kaufmann 1992b, Walde et al. 2007). Similar to the environmental temperatures in our study, nesting occurred when AT and ST were 22.0–26.3 °C and 21.5–25.5 °C, respectively, in Pennsylvania (Ernst 1986). Soil temperature is the most significant contributor to Wood Turtle nest-site selection, with nesting occurring on beaches with medium sand to larger grains (Hughes et al. 2009). In our population, preferred nesting beaches were primarily Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 525 composed of sand intermixed with pebbles and minimal vegetative cover, possibly chosen for their thermal characteristics. Wood turtles also have nested within natural erosion zones (Saumure et al. 2007), agricultural fields (Kaufmann 1992a), gravel pits (Walde et al. 2007), sandy roadsides (Quinn and Tate 1991), and sandpits with sparse vegetation (Tuttle and Carroll 2005). High percentages of Wood Turtle females returned to the same nesting areas annually (Walde et al. 2007) and would make several uncompleted nest holes (Harding and Bloomer 1979, Parren 2013), possibly to test the soil for suitability or to deter nest predators (Harding and Bloomer 1979). In our study, beaches were shared among multiple females, with several test nests observed. Depredated nests (indicated by shredded, scattered eggshells surrounding an excavated nest) were often found less than 0.5 m from each other, although we never observed predators. Marchand et al. (2002) found 22% of artificial nests to be disturbed within a week of placement; Procyon lotor (L.) (Raccoon) was the most common predator. Clumped nests and nests in agricultural or disturbed areas, like our study, were more often preyed upon than scattered nests or nests near roads or in manicured lawns (Marchand et al. 2002, Marchand and Litvaitis 2004). However, studies have not all reported obervations of mammalian predation of Wood Turtle (Parren 2013, Walde et al. 2007). In our study, of the 11 eggs laid, only 1 failed to develop and 7 emerged when we were not present. The 3 hatchlings seen emerging began trying to eat their eggshells soon thereafter. We speculate that this may be a need to absorb calcium to strengthen their shells, which are soft when the young hatch. Comparatively, clutch sizes varied from 5 to 11 individuals in New Jersey (Farrell and Graham 1991), 5 to 18 in Michigan (Harding and Bloomer 1979), and 5 to 20 in Québec (Walde et al. 2007). Our West Virginia hatchlings were larger than their New Hampshire counterparts, possibly suggesting that the trend of southern populations of turtles being larger than middle-of-the-range populations may begin during development in the egg. Active season. After spring mating and nesting, the turtles became more active and solitary. Wood Turtles in our population tended to walk along the river bottom rather than swim, noted also by Brewster and Brewster (1991), possibly providing greater traction when moving in the water. Water channels near Wood Turtle populations varied in width and tended to have sandy substrates with large scattered rocks and logs (Brewster and Brewster 1991, Greaves and Litzgus 2007). Stream width and depth in our study was larger than in Wisconsin (3–5 m wide and 0.3–1.5 m deep; Brewster and Brewster 1991) but similar to Canada (10–20 m wide and less than 2 m deep; Greaves and Litzgus 2007). In our West Virginia study, temperatures related to aquatic movement were similar to other regional temperatures, but with higher upper limits (air = 42 °C, water = 29 °C, and soil = 34 °C) related to our more southern latitude. Aquatic movement occurred in Pennsylvania during AT = 3.0–26.0 °C and WT = 6.0– 20.0 °C (Ernst 1986) and in New Jersey when AT = 3.6–24.8 °C and WT and ST both at 4.0–25.0 °C (Farrell and Graham 1991). The terrestrial period of our Wood Turtle population was consistent with observations of other populations in their geographic range, with higher upper limits in the south. Terrestrial activity was prominent by June in Pennsylvania (Ernst 1968, Northeastern Naturalist 526 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 Vol. 25, No. 4 Kaufmann 1992a) during AT = 14.0–32.0 °C and ST = 14.0–28.0 °C (Ernst 1986) and in New Jersey when AT = 11.0–27.4 °C and WT and ST both at 11.2–27.0 °C (Farrell and Graham 1991). During our high southern extremes, the turtles estivated, burrowing into vegetation, leaf litter, cow-hoof depressions, log jams, and thick mud to escape the heat, suggesting hot temperatures may be a limiting factor to Wood Turtles along the southern border of their geographic range (McCoard et al. 2016b). However, cover on land has been used also to escape a perceived threat (Parren 2013). During cooler temperatures, our turtles were often seen walking along deer trails, possibly because they were visually characteristic of dry streambeds (Tuttle and Carroll 2005, Yeomans 1995). Although positive geotaxis is beneficial to aquatic and semi-aquatic turtle species for finding water (DeRosa and Taylor 1980, Tuttle and Carroll 2005), movement both downhill and uphill along existing paths was observed in our study and in South Carolina (Yeomans 1995). Perhaps it may also be instinctual for Wood Turtles to follow trails; Tinklepaugh (1932) determined that Wood Turtles could find their way through mazes and Tuttle and Carroll (2005) observed that hatchlings followed each other’s trails. Travelling along existing paths may be less energetically costly for the turtles than forging new paths. Basking. Our Wood Turtles basked terrestrially at a 45° angle towards the sun primarily before noon with their heads and appendages extended. Similarly, they basked at stream edges in Québec (Arvisais et al. 2002, Saumure and Bider 1998), on banks and floodplains along streams before noon in Pennsylvania (Ernst 1986, Kaufmann 1992a), and in streambank depressions at 25–80° angles in New Jersey (Farrell and Graham 1991). In New Hampshire, hatchlings basked at 45°, front legs extended, after emerging from overnight locations (Tuttle and Carroll 2005). Basking occurred in our population during temperatures similar to those in other parts of the Wood Turtles’ range: AT =14.0–33.0 °C, WT = 12.0–24.0 °C, and ST = 14.0–32.0 °C in Pennsylvania (Ernst 1986); and AT = 4.0–29.4 °C and WT and ST both at 3.4–34.4 °C in New Jersey (Farrell and Graham 1991). Diet. Wood Turtle diet is fairly uniform across their range. In our West Virginia population, turtles were observed eating sooner, consistent with earlier emergence from hibernation, but terminating at the same time as for northern Wood Turtles. In Québec, early July through September was noted as a heavy feeding period (Arvisais et al. 2002). The omnivorous diet observed in our study is consistent with other diet observations (Castellano and Behler 2003, Compton et al. 2002, Farrell and Graham 1991, Strang 1983, Tuttle and Carroll 2005, Walde et al. 2003). Wood Turtles in our population, as in Pennsylvania, were seen worm-stomping (Kaufmann 1986). Dominance and aggression. The sequence of events in an aggressive male:male encounter observed during our study were similar to observations made by Kaufmann (1992b). A ranking system appears to exist in Wood Turtle populations based on age and mass, with higher percentages of aggressive, compared to non-aggressive, male:male encounters primarily occurring aquatically in autumn (Kaufmann 1992b). Harding and Bloomer (1979) observed dominance behavior in Northeastern Naturalist Vol. 25, No. 4 K.R.P. McCoard, N.S. McCoard, and J.T. Anderson 2018 527 captive Wood Turtles; males tended to be more dominant in aquatic situations and when of equal or larger size than females, but females were more dominant in terrestrial situations and when larger than males. Brumation. In our West Virginia population, Wood Turtles returned to the river in October and brumated communally from November to March in thick mud, root masses, or exposed on the riverbed. After a warm spell in late February to early March 2011, the female turtle that we found deceased during the brumation period was in a water depth (0.25 m) that we speculated to be too shallow to act as a buffer against the sudden, returning cold temperatures, which would suggest that Wood Turtles require a specific minimum depth to facilitate survival d uring brumation. In Pennsylvania, Wood Turtles brumated from late October to early April under overhanging streambanks, exposed tree roots, or in stream substrate (Kaufmann 1992a). In Québec, Wood Turtles begin to brumate in November in the riverbank or on the streambed (Arvisais et al. 2002, 2004). In Vermont, they brumated communally from November to March, using logs, rocks, leaves, silt, and small coves as cover and occasionally moving locations (Parren 2013). In Ontario, Wood Turtles selected mucky substrate over sandy substrate (Greaves and Litzgus 2008). Turtles in our study were occasionally found brumating under ice in calm stream reaches, a situation also observed in Algonquin Park, Canada (Quinn and Tate 1991) as well as Vermont (Parren 2013), but not Pennsylvania (Ernst 1986). In our study, the average brumation depth and average distance from shore was similar to observations in Ontario (depth = 0.91–1.00 m, mean distance from shore = 1.00–1.24 m; Greaves and Litzgus 2007, 2008) and Pennsylvania (depth = 1.0–2.3 m; Ernst 1986). Future research. Our natural history observations on a Wood Turtle population in West Virginia, near the southern extent of their geographic range, aid in filling data gaps on the species. This information is essential for conservation and management practices to be effective and successful for a species that is in decline. Further research is needed on nest predation, heat tolerance, and the impacts of agricultural practices on survival and juvenile recruitment of Wood Turtles within the southern portion of their range. Acknowledgments We captured the turtles under permits from the West Virginia Division of Natural Resources and the West Virginia University Animal Care and Use Committee, protocol 09-0408. We thank C.L. Pawlik, S. Selego, M. Jones, C. Concepcion, and L. Moon for assistance in the field. Previous drafts of this manuscript were reviewed by T.K. Pauley, D.J. Brown, A.A. Billings, P. Bohall-Wood, and E.D. Michael. Many landowners allowed us entrance onto their properties, and we thank them for the access. Funding for this project was provided by the Chesapeake Bay Conservation Innovation Grant Program (USDA), the National Fish and Wildlife Foundation, the National Oceanic and Atmospheric Administration, and the West Virginia University Natural History Museum. J.T. 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