Home Range and Habitat Selection in the Eastern Box Turtle (T )

Home , Eastern box turtle, Florida box turtle

Herpetological Conservation and Biology 10(1):99-111. Submitted: 24 October 2014; Accepted: 4 December 2014; Published: 14 June 2015. Home Range and Habitat Selection in the Eastern Box Turtle (Terrapene carolina carolina) in a Longleaf Pine (Pinus palustris)Reserve

Sasha E. Greenspan1,4, Erin P. Condon2 and Lora L. Smith3

1School of Marine and Tropical Biology, James Cook University, Douglas, Queensland 4811, Australia 2614 SW 27th Street, Gainesville, Florida 32607, USA 3Joseph W. Jones Ecological Research Center, 3988 Jones Center Drive, Newton, Georgia 39870, USA 4Corresponding author, e-mail: [email protected]

Abstract.—The Eastern Box Turtle (Terrapene carolina) is a species of conservation concern throughout North America, with recent population declines attributed primarily to habitat loss. The habitat requirements of this species in the southeastern United States have not been fully described. Our objectives were to estimate home range size and to describe habitat selection of Eastern Box Turtles (subspecies T. c. carolina) in a landscape dominated by Longleaf Pine (Pinus palustris) forest, the once dominant ecosystem in the Southeastern Coastal Plain. We conducted a radio-telemetry study of adult Eastern Box Turtles in a managed Longleaf Pine reserve in southwestern Georgia, USA. Home ranges expressed as 95% minimum convex polygons were 0.33–54.37 ha in size and averaged 10.33 ± 3.33 ha (SE). Turtles exhibited landscape-scale selection of pine-hardwood forests and hardwood forests. At a local scale, turtles used forbs more than bare ground, litter, and grass. Our study provides much- needed baseline information about home range size and habitat use of Eastern Box Turtles in the endangered Longleaf Pine ecosystem. Additional studies, particularly regarding use of disturbed habitats that are more characteristic of the modern southeastern landscape, would further clarify the status of Eastern Box Turtles in this region.

Key Words.—Georgia; kernel density estimate; minimum convex polygon; radio-telemetry.

Introduction such as expanding road networks leading to vehi- cle strikes, habitat degradation and loss, and in- Eastern Box Turtles (Terrapene carolina) oc- ﬂated populations of human-associated predators cur in the eastern and midwestern US, south- (Williams and Parker 1987; Hall et al. 1999), but ern Ontario, Canada, and northern Mexico (Carr commercial collection (Gibbons et al. 2000) and 1952; Dodd 2001). As a result of gradual, but infectious diseases (Feldman et al. 2006; John- at times pronounced, population declines over son et al. 2008) are also contributing factors. De- recent decades (e.g., Stickel 1978; Williams and creased adult survival, a hallmark of most docu- Parker 1987; Hall et al. 1999), the species is mented declines in Eastern Box Turtles, is espe- listed in CITES Appendix II as vulnerable in the cially signiﬁcant from a conservation perspective District of Columbia, Massachusetts, New York, (Hall et al. 1999). The delayed maturity and slow and Pennsylvania, and as critically imperiled in rate of population growth that are typical of this Maine and New Hampshire (NatureServe. 2013. species lead to slow recovery from losses (Hall NatureServe Explorer: an online encyclopedia et al. 1999). The conservation status of the East- of life. Available from http://www.natureserve. ern Box Turtle warrants a more comprehensive org/explorer [Accessed 1 July 2014]). Most de- understanding of its natural history and behavior, clines are linked to land use changes by humans, key components of which are area requirements

Copyright c 2015. Sasha E. Greenspan. All Rights Reserved. 99 Greenspan et al.—-Eastern Box Turltes in Longleaf pines. and habitat selection (Stickel 1950; Conner and this study were to describe home range size of Godbois 2003). With the exception of extensive Eastern Box Turtles (subspecies T. c. carolina) data for an isolated population on Egmont Key and to examine habitat selection at multiple spa- in Tampa Bay, Florida (Dodd et al. 1994; Dodd tial scales in a Longleaf Pine reserve managed 2001; Jennings 2003), these attributes are largely with prescribed fire. unspecified for populations south of Maryland (Stickel 1950, 1989; Dodd 2001). Materials and Methods The home range of an animal refers to the area routinely used for daily activities including Study site.—We conducted our study at resting, foraging, thermoregulation, mating, and Ichauway, a privately owned reserve and the nesting (Burt 1943; Stickel 1950, 1989; Dodd site of the Joseph W. Jones Ecological Research 2001). Within a home range, most individuals Center in Baker County, Georgia, USA (Fig. use certain core areas more than others, presum- 1). Ichauway is dominated by second growth ably because resources are not evenly distributed Longleaf Pine forest and has been intensively across the landscape (Worton 1989; Chamber- managed with prescribed fire since the early lain et al. 2003). Research throughout the range 1900s. The 11,769-ha property also contains of the Eastern Box Turtle suggests that home other pines (e.g., Slash Pine, Pinus elliottii, ranges are typically from 1–5 ha in size (Dodd Loblolly Pine, P. taeda, and Short-leaf Pine, P. 2001) but can reach about 20–25 ha (Schwartz echinata), hardwoods (typically oaks, Quercus et al. 1984; Ernst and Lovich 2009). Variation in spp.), wildlife food plots for the Northern home range estimates may be due to geographic Bobwhite (Colinus virginianus) and other location and habitat quality (Kaufmann 1995; game species (Boring 2001), and numerous Nieuwolt 1996; Arvisais et al. 2002; Chamber- semi-permanent to extremely ephemeral wet- lain et al. 2003). Thus, it is important to estimate lands (Drew et al. 1998; Kirkman et al. 1998; home range across a variety of habitat types and Fig. 1). Two large streams, Flint River and geographic regions, especially in areas where pre- Ichawaynochaway Creek, and several paved vious work has been limited. highways intersect the property. Center-pivot In the eastern portion of its range, the Eastern agriculture is the primary land use surrounding Box Turtle is generally associated with mesic the reserve. forests, which meet the thermal and hydric requirements of the species (Reagan 1974; Dodd Radio-telemetry.—We collected Eastern Box 2001). Longleaf Pine (Pinus palustris) forest was Turtles opportunistically throughout Ichauway once the dominant forest type in the Southeastern between August 2003 and March 2004. For each Coastal Plain but has been reduced to < 3% of turtle, we measured body mass and straight-line its historic range (Frost 1993). In light of this carapace length (CL) and determined sex based dramatic loss, it is important to determine the de- on the degree of plastron concavity and eye color gree to which Eastern Box Turtles are associated (Nichols 1939). We individually marked turtles with Longleaf Pine forests versus other available by notching the marginal scutes with a triangu- habitats in the southeastern United States. This is lar file (Cagle 1939). We outfitted 23 adult tur- especially necessary considering that prescribed tles (> 100 mm CL) with SM-1H radio trans- fire is an essential tool in restoration and man- mitters weighing 26 g (AVM Instrument Co. agement of remaining Longleaf Pine stands, but Ltd., Livermore, California, USA). Transmitters whether fire provides favorable habitat character- were affixed to the costal scutes of the carapace istics for Eastern Box Turtles in the southeastern with epoxy putty (Oatey Supply Chain Services, region is not well documented. The objectives of Cleveland, Ohio, USA). The weight of the radio

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Figure 1. Study area for home range and habitat selection study of Terrapene carolina carolina at Ichauway reserve, Baker County, Georgia, USA.

101 Greenspan et al.—-Eastern Box Turltes in Longleaf pines. transmitter and fixative averaged < 40 g (approx- Longleaf Pine or Slash Pine. Aquatic areas were imately 10% of the body mass of the turtle). forested and non-forested isolated wetlands, the We released radio-tagged turtles at the point Ichawaynochaway Creek, and the Flint River. of capture and located each turtle at least once Shrub/scrub included abandoned clearcuts and per week during the active season (March- agricultural fields, failed pine plantations, and November) and once every two weeks during hardwood scrub. Developed areas were roads, the inactive season (December-March) between houses and sheds. September 2003 and October 2004. We tracked For each turtle, we calculated two estimates turtles using an R1000 radio-telemetry receiver of home range size, 95% minimum convex poly- (Communications Specialists Inc., Orange, gon (MCP; Mohr 1947) and 95% kernel den- California, USA) and a folding Yagi 6-element sity estimate (KDE; Worton 1989), and one es- antenna (Wildlife Materials Inc., Murphysboro, timate of core area size (50% KDE) using the Illinois, USA). We radio-tracked turtles until Home Range Tools extension (HRT; Rodgers signals were lost or the study ended (October et al. 2007) in ArcMap. We selected smoothing 2004). We recorded turtle locations using either factors (h) that generated 95% KDEs similar in a Trimble GeoExplorer 3 (accurate to within 10 area to the 95% MCPs (Row and Blouin-Demers m) or a TrimbleARC1 (accurate to within 1 m; 2006). We used linear regression (PROC REG) Trimble Navigation Systems Ltd., Sunnyvale, to determine whether home range size estimates California, USA). We removed transmitters at were related to length of the tracking period the completion of the study. (days) or number of locations. We defined the study area as a 2,046-ha area Data analysis.—We performed statistical anal- encompassed by 500-m circular buffers around yses with SAS 9.3 software (SAS Institute Inc., each MCP. When buffers of individual turtles Cary, North Carolina, USA). We set a signifi- overlapped, we used the Dissolve Tool to cre- cance level of α = 0.05. We performed spatial ate a single, continuous polygon (Johnson l980; analyses with ArcMap 9.3.1 software (ESRI, Red- Fig. 1). A small portion of the study area fell out- lands, California, USA). We used an existing ge- side of Ichauway in an area for which we had no ographic information system delineating habitat habitat data. However, none of the turtles moved types for Ichauway, produced by screen digitiz- outside of Ichauway, indicating that the compo- ing and ground-truthing 2006 color infrared dig- sition of the study area with habitat data was ital aerial photography at 1-ft pixel resolution. representative of habitat available to the turtles. Habitat types were pine forest, hardwood forest, We used compositional analysis, a test of pro- pine-hardwood forest, agricultural, pine planta- portional habitat use relative to availability, to tion, aquatic, shrub/scrub, and developed. Pine assess habitat selection by turtles, both with and forest referred to mixed-age stands with > 90% without sex-specific comparisons (Aebischer et Longleaf Pine (dominant across 90% of this habi- al. 1993). To assess habitat selection at the lo- tat type), Slash Pine, Loblolly Pine, or Shortleaf cal scale, we compared habitat composition at Pine. Hardwood forest contained > 90% hard- unique turtle locations (i.e., telemetry locations woods including Red Oak (Q. falcata), Live Oak in which the turtle had moved > 5 m from its (Q. virginiana), Laurel Oak (Q. laurifolia), or preceding location) to habitat composition of the Water Oak (Q. nigra). Pine-hardwood forest was home ranges (95% KDE and 95% MCP), and composed of 20–80% pines or hardwoods. Agri- habitat composition of the core areas (50% KDE) cultural land comprised wildlife food plots and to that of the home ranges (95% KDE). To as- other small croplands. Pine plantation consisted sess habitat selection at the landscape scale, we of newly planted to mature even-aged stands of compared both habitat composition of the home

102 Herpetological Conservation and Biology ranges (95% KDE and 95% MCP) and of the core eraged 10.33 ± 3.33 ha. The average MCP for areas (50% KDE) to that of the study area (John- males (11.70 ± 5.75 ha) was larger than that son 1980). We performed multivariate analyses for females (8.33 ± 5.68 ha). KDE home ranges of variance (MANOVA) of log-ratio transformed were similar in area to MCPs, as we designed. data (PROC GLM). When a MANOVA indicated Core areas were 0.09–10.44 ha and averaged 2.08 selection, we identified the habitat types selected ± 0.62 ha. The average core area for males (2.33 by turtles with a ranking matrix of t-tests (PROC ± 1.10 ha) was also larger than that for females MEANS). (1.60 ± 0.99 ha; Appendix 1). Home range size We assessed groundcover preferences with use- estimates were not significantly related to length availability analyses (Neu et al. 1974). Here, we of time tracked (r2 = 0.021, P = 0.554) or number define use as the groundcover type at unique tur- of tracking events (r2 = 0.139, P = 0.116). The tle locations, i.e., whenever a turtle moved > relatively high degree of variation in MCPs was 5 m from its preceding location, we recorded due to two individual turtles with exceptionally the groundcover type on which it was presently large home ranges (male 1135, MCP = 54.37 sitting. Groundcover types were bare ground ha and female 1142, MCP 42.03 ha, Appendix (no vegetation), litter (leaves and pine nee- 1). For comparison, the average MCP excluding dles), grasses (predominantly Wiregrass, Aris- these two individuals was 5.87 ± 1.36 ha. tida stricta, and Broomsedge, Andropogon spp.) Habitat use did not differ between female and and forbs (predominantly oak seedlings, Quer- male turtles (F7,12 = 0.270, P = 0.950), so we cus spp., Silkgrass, Pityopsis graminifolia, black- report results of models without a sex compo- berry, Rubus spp., and Poison Oak, Toxicoden- nent. At the landscape scale, habitat composition dron radicans). For analyses, data were summa- of the study area differed from that of core ar- rized as percentage of all unique locations within eas (F7,12 = 9.94, P < 0.001) and home ranges each of the different groundcover types. We esti- (KDE: F7,12 = 6.99, P = 0.002; MCP: F7,12 = mated groundcover availability with an existing 10.48, P < 0.001). Specifically, the proportion data set for Ichauway that included percentages of mixed pine-hardwood and hardwood forest of groundcover types within 442 randomly placed in home ranges and core areas was greater than 1-m2 quadrats (Miller 2008). We used chi-square expected based on availability, whereas the pro- statistics (PROC FREQ) to test the hypothesis portions of agricultural land, aquatic habitats, de- that groundcover types were used in proportion veloped areas, pine forests, pine plantations, and to their availability. If the null hypothesis was scrub-shrub habitat in home ranges were less than rejected (α < 0.05), we employed confidence in- expected (Fig. 2). At the local scale, habitat com- tervals following methods in Neu et al. (1974) to position of home ranges did not differ from that of determine which groundcover types were used in core areas (KDE: F7,12 = 0.96, P = 0.498) or tur- proportions greater or less than availability. tle locations (MCP: F7,12 = 1.08, P = 0.431; KDE: F7,12 = 2.10, P = 0.123). Eastern box turtle use Results of groundcover differed from random ( χ2 = 32.9, df = 3, P < 0.001); turtles preferred groundcover We radio-tracked 19 turtles (seven female, nine comprised of forbs and avoided bare ground and male, three of undetermined sex) for at least 4 litter (Fig. 3). mo of the active season. We excluded four individuals that were lost within 4 mo of track- Discussion ing from analyses. We tracked individuals from 35–90 times over 174–401 d (x¯ = 326 d; Ap- To our knowledge, this is the first account of pendix 1). MCPs were 0.33–54.37 ha and av- home range size and habitat selection of Eastern

103 Greenspan et al.—-Eastern Box Turltes in Longleaf pines.

Figure 2. Average proportional habitat use (95% MCP) by Terrapene carolina carolina relative to availability (study area) at Ichauway reserve, Baker County, Georgia, USA. Habitat compositions of KDEs were similar to MCPs and are not shown.

Figure 3. Average proportional groundcover use by Terrapene carolina carolina relative to availability at Ichauway reserve, Baker County, Georgia, USA.

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Box Turtles in a Longleaf Pine landscape matrix. suitable habitat to ensure that large home ranges Longleaf Pine-dominated forests at Ichauway and can be accommodated when they are essential for throughout the Southeastern Coastal Plain are survival and reproduction. managed with frequent prescribed fire. Studies in Our compositional analysis revealed habitat other regions have reported injuries and mortality selection at the landscape scale but not within of box turtles from fire (Babbitt and Babbitt 1951; home ranges, underscoring the value of a two- Platt et al. 2010). Our findings indicate, however, stage approach to analyses of habitat use (Ae- that despite risk to individuals from occasional bischer et al. 2003). Specifically, the turtles that fires, fire-maintained forests may offer important we tracked favored pine-hardwood forest and benefits to Eastern Box Turtle populations in the hardwood forest in our study area. On Ichauway, southeastern United States. pine-hardwood forest is found on a wide soil Home range sizes of Eastern Box Turtles on moisture gradient from upland sand ridges to the Ichauway reserve were highly variable as wetland edges but is generally characterized as has been reported for the species in other habi- a mesic habitat with a significant component tats (Dodd 2001). Previously reported average of Longleaf Pine (Kirkman et al. 2001). Hard- MCPs in the southeastern US are 1.88 ha (Ten- wood forests on Ichauway generally occur in nessee: Donaldson and Echternacht 2005), 2.68 proximity to streams (Fig. 1) on more mesic ha (North Carolina: Kapfer et al. 2013), and 6.45 soils (Jacqmain et al. 1999; Goebel et al. 2001). ha (North Carolina: Hester et al. 2008). Average The association between Eastern Box Turtles and home range size in our study was comparatively mesic forests is well supported by previous work high (10.33 ha) owing to one male and one fe- (reviewed by Dodd 2001) and is linked to the male with home ranges > 40 ha. For the male species’ thermal and hydric requirements (Rea- (#1135; Appendix 1), tracking coincided with gan 1974; Rossell et al. 2006). Mesic forests typ- two long movements across and returning across ically have dense tree canopies. These canopies a two-lane highway in April (1,913 m over 8 d in can moderate temperatures and retain understory one direction and 1,757 m over 2 d on the return). humidity but still permit intermittent sunlight pen- The female (#1142) moved at least 553 m over etration and basking opportunities (Dodd 2001). 3 d in July (184.4 m/d). Box turtles in Georgia On Ichauway and throughout the Southeastern mate from April to June and nest in late spring or Coastal Plain, frequent fire leads to similarities in summer (Cash and Gibbons 2008); thus, it is pos- floral attributes between pine-hardwood forests sible that these extended movements over a short and Longleaf Pine savanna. Similarities include period of time were related to mate searching and diverse native groundcover (Lemon 1949; Walker nesting (Stickel 1950; Iglay et al. 2007). Exclud- and Peet 1984; Kirkman et al. 2001) and high ing these two individuals, our estimate of MCP faunal diversity (Guyer and Bailey 1993), with home range size was 5.87 ha, which is closer to the oak component adding canopy cover, verti- that reported in surveys elsewhere in the south- cal structure, seedlings, and coarse woody debris. eastern US (Donaldson and Echternacht 2005, Although the ecological role of oaks in Longleaf Hester et al. 2008, Kapfer et al. 2013). Addi- Pine ecosystems is heavily debated, Hiers et al. tional long-term research is needed to more fully (2014) contend that increased canopy closure by predict the spatial requirements of the species pyrophytic oaks (i.e., oak species such as Q. fal- at the regional level and in response to land use cata that are associated with fire-prone uplands) change from urban and agricultural development. typically does not significantly affect understory However, our home range findings highlight the diversity at the stand scale. They also suggest importance in conservation planning of protect- that presence of pyrophytic oaks may facilitate ing wildlife corridors and contiguous tracts of increased arthropod diversity and biomass, thus

105 Greenspan et al.—-Eastern Box Turltes in Longleaf pines. potentially increasing habitat quality (e.g., for- for assistance with GIS. Mike Conner and Jen age) for some species, including the Eastern Box Howze also assisted with data analyses and re- Turtle. At a fine scale, turtles in our study pre- viewed an earlier draft of this manuscript. We ferred patches of forbs and avoided bare ground, also thank Jones Center staff for collecting turtles litter, and grass substrates. Additional research for this study. This paper was written in memory is needed to determine whether this groundcover of Dr. Larry Galblum. provides more suitable foraging and thermoregu- latory environments than other available ground- Literature Cited cover types. If this is the case, it might explain our observation that turtles avoided pine planta- Aebischer, N.J., P.A. Robertson, and R.E. Ken- tions and agricultural areas, which lack ground- ward. 1993. Compositional analysis of habitat cover vegetation. use from animal radiotracking data. Ecology Our results contribute new information to the 74:1313–1325. body of literature on the habitat requirements of this species in the southeastern United States Arvisais, M., J.C. Borgeois, E. Levesque, C. and shed light on the value of ecological aware- Daigle, D. Masse, and J. Jutras. 2002. Home ness in regional forest management. Specifically, range and movements of a Wood Turtle (Clem- although removal of hardwoods is typically a mys insculpta) population at the northern limit major element of Longleaf Pine forest manage- of its range. Canadian Journal of Zoology ment, our study supports the view that a modest 80:402–408. upland-associated oak component can increase Babbitt, L.H., and C.H. Babbitt. 1951. A her- the ecological value of Longleaf Pine forests petological study of burned-over areas in Dade when combined with prescribed fire (Hiers et County, Florida. Copeia 1951:79. al. 2014). Knowledge of home range size and habitat selection of Eastern Box Turtles is critical Boring, L.R. 2001. The Joseph W. Jones Eco- for establishing habitat conservation and restora- logical Research Center: co-directed applied tion priorities, especially considering continuing and basic research in the private sector. Pp. loss of forested habitats to urban and agricul- 233–258 In Holistic Science: The Evolution of tural development throughout the range of this the Georgia Institute of Ecology (1940–2000). species and growing concern for infectious dis- Barrett, G.W., and T.L. Barrett (Eds.). Taylor eases (Feldman et al. 2006; Johnson et al. 2008). & Francis, New York, New York, USA. Given the dominance on the southeastern landscape of large-scale agriculture and urban/ subur- Burt, W.H. 1943. Territoriality and home range ban development, we encourage future research concepts as applied to mammals. Journal of on the habitat preferences of Eastern Box Turtles Mammalogy 2:346–352. in non-forested sites. Cagle, F.R. 1939. A system for marking Acknowledgements.—This study was partially turtles for future identification. Copeia funded by the University of Florida and the 1939:170–172. Joseph W. Jones Ecological Research Center. Work was conducted under Georgia Department Carr, A. 1952. Handbook of Turtles. Comstock of Natural Resources Scientific Collecting Per- Publishing Associates, Ithaca, New York, USA. mits 29-WMB-03-142 and 29-WMB-04-84. We thank Dick Franz and George Tanner from the Chamberlain, M.J., B.D. Leopold, and L.M. Con- University of Florida for guidance throughout the ner. 2003. Space use, movements, and habitat field study. We thank Jean Brock and Jen Howze

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selection of adult Bobcats (Lynx rufus) in cen- Frost, C.C. 1993. Four centuries of changing tral Mississippi. American Midland Naturalist landscape patterns in the Longleaf Pine ecosys- 149:395–405. tem. Proceedings of the Tall Timbers Fire Ecol- ogy Conference 18:17–43. Cash, W.B., and J.W. Gibbons. 2008. Eastern box turtle. Pp.407–499 In Amphibians and Gibbons, J.W., D.E. Scott, T.J. Ryan, K.A. Reptiles of Georgia. Jensen, J.B., C.D. Camp, Buhlmann, T.D. Tuberville, B.S. Metts, J.L. W. Gibbons, and M.J. Elliott (Eds). University Greene, T. Mills, Y. Leiden, S. Poppy, et al. of Georgia Press, Athens, USA. 2000. The global decline of reptiles, déjà vu amphibians. BioScience 50:653–666. Conner, L.M., and I.A. Godbois. 2003. Habitat associated with daytime refugia of Fox Squir- Goebel, C.P., B.J. Palik, L.K. Kirkman, M.B. rels in a Longleaf Pine forest. American Mid- Drew, L. West, and D.C. Pederson. 2001. For- land Naturalist 150:123–129. est ecosystems of a Lower Gulf Coastal Plain landscape: multifactor classification and anal- Dodd, C.K. 2001. North American Box Turtles: ysis. Journal of the Torrey Botanical Society a Natural History. The University of Oklahoma 128:47–75. Press, Norman, Oklahoma, USA. Guyer, C., and M.A. Bailey. 1993. Amphibians Dodd, C.K., R. Franz, and L.L. Smith. 1994. Ac- and reptiles of longleaf pine communities. Pro- tivity patterns and habitat use of box turtles ceedings of the Tall Timbers Fire Ecology Con- (Terrapene carolina bauri) on a Florida island, ference 18:139–159. with recommendations for management. Che- lonian Conservation and Biology 1:97–106. Hall, R. J., P.F.P. Henry, and C.M. Bunck. 1999. Fifty-year trends in a box turtle popu- Donaldson, B.M., and A.C. Echternecht. 2005. lation in Maryland. Biological Conservation Aquatic habitat use relative to home range 88:165–172. and seasonal movement of Eastern Box Tur- tles (Terrapene carolina carolina: Emydidae) Hester, J.M., S.J. Price, and M.E. Dorcas. 2008. in Eastern Tennessee. Journal of Herpetology Effects of relocation on movements and home 39:278–284. ranges of Eastern Box Turtles. Journal of Wildlife Management 72:772–777. Drew, M.B., L.K. Kirkman, and A.K. Ghol- son, Jr. 1998. The vascular flora of Ichauway, Hiers, J.K., J.R. Walters, R.J. Mitchell, J.M. Baker County, Georgia: a remnant Longleaf Varner, and L.M. Conner. 2014. Ecological Pine/Wiregrass ecosystem. Castanea 63:1–24. value of retaining pyrophytic oaks in Longleaf Ernst, C.H., and J.E. Lovich. 2009. Turtles of the Pine ecosystems. Journal of Wildlife Manage- United States and Canada. The Johns Hopkins ment 78:383–393. University Press, Baltimore, Maryland, USA. Iglay, R.B., J.L. Bowman, and N.H. Nazdrowicz. Feldman, S.H., J. Wimsatt, R.E. Marchang, A.J. 2007. Eastern Box Turtle (Terrapene carolina Johnson, W. Brown, J.C. Mitchell, and J.M. carolina) movements in a fragmented land- Sleeman. 2006. A novel Mycoplasma detected scape. Journal of Herpetology 41:102–106. in association with upper respiratory disease Jacqmain, E.I., R.H. Jones, and R.J. Mitchell. syndrome in free-ranging Eastern Box Turtles 1999. Influences of frequent cool season burn- (Terrapene carolina carolina) in Virginia. Jour- ing across a soil moisture gradient on oak com- nal of Wildlife Diseases 42:279–289.

107 Greenspan et al.—-Eastern Box Turltes in Longleaf pines.

munity structure in longleaf pine ecosystems Miller, G.J. 2008. Home range size, habitat as- The American Midland Naturalist 141:85–100. sociations and refuge use of the Florida Pine snake, Pituophis melanoleucus mugitus, in Jennings, A.H. 2003. Daily and seasonal activity southwest Georgia, USA. M.S. Thesis, Uni- patterns and movements of juvenile box tur- versity of Florida, Gainesville. 72 pp. tles (Terrapene carolina bauri) on Egmont Key, Florida. Chelonian Conservation and Biology Mohr, C.O. 1947. Table of equivalent popula- 4:578–587. tions of North American small mammals. The American Midland Naturalist 37:223–249. Johnson, A.J., A.P. Pessier, J.F. Wellehan, A. Childress, T.M. Norton, N.L. Stedman, D.C. Neu, C.W., C.R. Byers, and J.M. Peek. Bloom, W. Belzer, V.R. Titus, R. Wagner, et 1974. A technique for analysis of utilization- al. 2008. Ranavirus infection of free-ranging availability data. Journal of Wildlife Manage- and captive box turtles and tortoises in the ment 38:541–545. United States. Journal of Wildlife Diseases Nichols J.T. 1939. Data on size, growth and age 44:851–863. in the box turtle, Terrapene carolina. Copeia Johnson, D.H. 1980. Comparison of usage and 1939:14–20. availability measurements for evaluating re- Nieuwolt, P.M. 1996. Movement, activity, and source preference. Ecology 61:65–71. microhabitat selection in the Western Box Tur- tle, Terrapene ornata luteola, in New Mexico. Kapfer, J.M., D.J. Muñoz, J.D. Groves, and R.W. Herpetologica 52:487–495. Kirk. 2013. Home range and habitat preferences of Eastern Box Turtles (Terrapene car- Platt, S.G., H. Liu, and C.K. Borg. 2010. Fire olina Linnaeus, 1758) in the Piedmont Ecolog- ecology of the Florida Box Turtle (Terrapene ical Province of North Carolina (USA). Her- carolina bauri Taylor) in pine rockland forests petology Notes 6:251–260. of the lower Florida Keys. Natural Areas Jour- nal 30:254–260. Kaufmann, J.H. 1995. Home ranges and movements of Wood Turtles, Clemmys insculpta, in Reagan, D.P. 1974. Habitat selection in the central Pennsylvania. Copeia 1995:22–27. Three-toed Box Turtle, Terrapene carolina tri- unguis. Copeia 1974:512–527. Kirkman, L.K., M.B. Drew, L.T. West, and E.R. Blood. 1998. Ecotone characterization be- Rodgers, A.R., A.P. Carr, H.L. Beyer, L. Smith, tween upland Longleaf Pine / Wiregrass stands and J.G. Kie. 2007. HRT: Home Range Tools and seasonally-ponded isolated wetlands. Wet- for ArcGIS. Ontario Ministry of Natural Re- lands 18:346–364. sources, Centre for Northern Forest Ecosystem Research, Thunder Bay, Ontario, Canada. Kirkman, L.K., R.J. Mitchell, R.C. Helton, and M.B. Drew. 2001. Productivity and species Rossell, Jr., C.R., I.M. Rossell, and S. Patch. richness across an environmental gradient in 2006. Microhabitat selection by Eastern Box a ﬁre-dependent ecosystem. American Journal Turtles (Terrapene c. carolina) in a North Car- of Botany 88:2119–2128. olina mountain wetland. Journal of Herpetol- ogy 40:280–284. Lemon, P.C. 1949. Successional responses of herbs in the Longleaf-Slash Pine forest after Row, J.R., and G. Blouin-Demers. 2006. Kernels ﬁre. Ecology 30:135–145. are not accurate estimators of home range size for herpetofauna. Copeia 2006:797–802.

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Schwartz, E.R., C.W. Schwartz, and A.R. Kiester. 1984. The Three-toed Box Turtle in Central Missouri, Part II: A Nineteen Year Study of Home Range, Movements and Population. Mis- souri Department of Conservation Terrestrial Series 12.

Stickel, L.F. 1950. Populations and home range relationships of the box turtle, Terrapene c. carolina (Linnaeus). Ecological Monographs 20:351–378.

Stickel L.F. 1978. Changes in a box turtle population during three decades. Copeia 1978:221– 225.

Stickel, L.F. 1989. Home range behavior among box turtles (Terrapene c. carolina) of a bottom- land forest in Maryland. Journal of Herpetol- ogy 23:40–44.

Walker, J., and R.K. Peet. 1984. Composition and species diversity of pine-wiregrass savan- nas of the Green Swamp, North Carolina. Veg- etatio 55:163–179.

Williams, Jr., E.C., and W.S. Parker. 1987. A long-term study of a box turtle (Terrapene carolina) population at Allee Memorial Woods, Indiana, with emphasis on survivorship. Her- petologica 43:328–335.

Worton, B.J. 1989. Kernel methods for esti- mating utilization distribution in home-range studies. Ecology 70:164–168.

109 Greenspan et al.—-Eastern Box Turltes in Longleaf pines. T est siae KE,BkrCut,Goga S,20–04 F 2003–2004. USA, radio-tracked Georgia, 19 County, for Baker ha) (KDE), (in estimates size density range home and period, tracking vrl 2 0 4361.31.02.08 1.60 2.33 10.30 (5.72) 11.70 8.28 10.33 11.70 3.11 1.78 8.33 326 2.37 0.28 0.49 12.04 8.85 0.25 11.56 3.13 1.56 34 12.25 3.81 3.05 0.80 9.04 1.49 11.38 10.44 1.24 13.44 1.54 0.52 20.75 0.37 343 3.11 3.69 0.09 371 1.47 53.99 0.71 257 13.26 5.36 20.48 174 2.00 7.41 1.51 1.78 3.52 339 32 407 0.36 0.65 54.37 335 38 3.14 0.22 5.38 350 33 2.12 336 41.62 373 26 1.43 7.13 371 46 0.33 127 3.37 77 240 35 3.01 0.87 73 478 391 33 42.02 65 401 126 29 Overall 7.30 322 35 32 3.34 174 317 70 18 area) 0.87 340 (core 330 74 132 KDE 50% 45 Males 350 375 68 50 330 63 390 26 Females 365 71 107 296 27 (SE) KDE Means 397 46 115 95% 428 22 85 124 450 35 (U) 1141 90 415 127 41 (U) 1137 76 MCP 359 109 37 (U) 1125 35 95% 385 132 41 (M) 1150 60 303 132 (M) 1146 Tracked 69 329 132 (M) 1144 77 491 132 (M) Days 1143 71 Locations 390 127 (M) 1138 78 560 Unique 119 No. (M) 1136 610 127 locations (M) 1135 360 130 Telemetry (M) 1131 570 122 (M) 1124 368 140 (F) 3000 140 (F) (g) 1151 Mass 123 (F) 1147 144 (F) 1142 122 (F) 1139 (mm) (F) 1133 CL (F) 1132 (gender) ID Turtle able A1. oysz tcpue(CL capture at size Body 2 2)()(5 33)(.0 (0.62) (1.10) (0.99) (3.30) (5.72) (5.62) (3.33) (5.75) (5.68) (15) (2) (21) (18) (2) (40) (3) (4) = aaaelnt) ubro ie utewsrdotakd ubro nqetakn oain,lnt of length locations, tracking unique of number radio-tracked, was turtle a times of number length), carapace erpn aoiacarolina carolina Terrapene = eae M female; = ae U male; = sn iiu ovxplgn MP n kernel and (MCP) polygons convex minimum using neemndsex. undetermined

110 Herpetological Conservation and Biology

Sasha E. Greenspan received a B.A. in environmental Erin P. Condon received her undergraduate degree in science and anthropology from Wesleyan University and Ecology and Evolutionary Biology from Princeton Uni- an M.S. in ecology and environmental science from the versity and attended graduate school in the Department of University of Maine. She is currently working toward a Wildlife Ecology and Conservation at the University of Ph.D. in tropical ecology at James Cook University in Florida from 2002–2004. Erin received J.D. and LL.M. Queensland, Australia. Her research and career interests degrees from the Frederic G. Levin College of Law and include amphibian and reptile ecology and conservation, is the former Executive Director of Florida Defenders tropical and disease ecology, One Health, and biologi- of the Environment in Gainesville, Florida, USA. (Pho- cal ﬁeld station operations. (Photographed by Stephen tographed by Sharon Rush). Harris).

Lora L. Smith is an Associate Scientist at the Joseph W. Jones Ecological Research Center in southwestern Geor- gia. Her work at the Center includes a long term study of the eﬀects of mammalian predators on Gopher Tortoises (Gopherus polyphemus), upland snake ecology, and habitat predictors of pond-breeding amphibians. Her research interests also include the eﬀects of habitat restoration and management on amphibian and reptiles. (Photographed by Miguel Pedrono).

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