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, Australia2614 SW 27th Street, Gainesville, Florida 32607, USA

3Joseph W. Jones Ecological Research Center, 3988 Jones Center Drive, Newton, Georgia 39870, USA4Corresponding author, e-mail: sasha.greenspan@gmail.com

Abstract.—The Eastern Box Turtle (Terrapene carolina) is a species of conservation concern through-out North America, with recent population declines attributed primarily to habitat loss. The habitatrequirements of this species in the southeastern United States have not been fully described. Our ob-jectives 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 oncedominant ecosystem in the Southeastern Coastal Plain. We conducted a radio-telemetry study of adultEastern Box Turtles in a managed Longleaf Pine reserve in southwestern Georgia, USA. Home rangesexpressed 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. Ata 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 theendangered Longleaf Pine ecosystem. Additional studies, particularly regarding use of disturbed habi-tats that are more characteristic of the modern southeastern landscape, would further clarify the statusof Eastern Box Turtles in this region.

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

Introduction

Eastern Box Turtles (Terrapene carolina) oc-cur in the eastern and midwestern US, south-ern Ontario, Canada, and northern Mexico (Carr1952; Dodd 2001). As a result of gradual, butat times pronounced, population declines overrecent decades (e.g., Stickel 1978; Williams andParker 1987; Hall et al. 1999), the species islisted in CITES Appendix II as vulnerable in theDistrict of Columbia, Massachusetts, New York,and Pennsylvania, and as critically imperiled inMaine and New Hampshire (NatureServe. 2013.NatureServe Explorer: an online encyclopediaof life. Available from http://www.natureserve.org/explorer [Accessed 1 July 2014]). Most de-clines are linked to land use changes by humans,

such as expanding road networks leading to vehi-cle strikes, habitat degradation and loss, and in-flated populations of human-associated predators(Williams and Parker 1987; Hall et al. 1999), butcommercial collection (Gibbons et al. 2000) andinfectious diseases (Feldman et al. 2006; John-son et al. 2008) are also contributing factors. De-creased adult survival, a hallmark of most docu-mented declines in Eastern Box Turtles, is espe-cially significant from a conservation perspective(Hall et al. 1999). The delayed maturity and slowrate of population growth that are typical of thisspecies lead to slow recovery from losses (Hallet al. 1999). The conservation status of the East-ern Box Turtle warrants a more comprehensiveunderstanding of its natural history and behavior,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 andGodbois 2003). With the exception of extensivedata for an isolated population on Egmont Keyin Tampa Bay, Florida (Dodd et al. 1994; Dodd2001; Jennings 2003), these attributes are largelyunspecified for populations south of Maryland(Stickel 1950, 1989; Dodd 2001).

The home range of an animal refers to thearea routinely used for daily activities includingresting, foraging, thermoregulation, mating, andnesting (Burt 1943; Stickel 1950, 1989; Dodd2001). Within a home range, most individualsuse certain core areas more than others, presum-ably because resources are not evenly distributedacross the landscape (Worton 1989; Chamber-lain et al. 2003). Research throughout the rangeof the Eastern Box Turtle suggests that homeranges are typically from 1–5 ha in size (Dodd2001) but can reach about 20–25 ha (Schwartzet al. 1984; Ernst and Lovich 2009). Variation inhome range estimates may be due to geographiclocation and habitat quality (Kaufmann 1995;Nieuwolt 1996; Arvisais et al. 2002; Chamber-lain et al. 2003). Thus, it is important to estimatehome range across a variety of habitat types andgeographic regions, especially in areas where pre-vious work has been limited.

In the eastern portion of its range, the EasternBox Turtle is generally associated with mesicforests, which meet the thermal and hydric re-quirements of the species (Reagan 1974; Dodd2001). Longleaf Pine (Pinus palustris) forest wasonce the dominant forest type in the SoutheasternCoastal Plain but has been reduced to < 3% ofits historic range (Frost 1993). In light of thisdramatic loss, it is important to determine the de-gree to which Eastern Box Turtles are associatedwith Longleaf Pine forests versus other availablehabitats in the southeastern United States. This isespecially necessary considering that prescribedfire is an essential tool in restoration and man-agement of remaining Longleaf Pine stands, butwhether fire provides favorable habitat character-istics for Eastern Box Turtles in the southeasternregion is not well documented. The objectives of

this study were to describe home range size ofEastern Box Turtles (subspecies T. c. carolina)and to examine habitat selection at multiple spa-tial scales in a Longleaf Pine reserve managedwith prescribed fire.

Materials andMethods

Study site.—We conducted our study atIchauway, a privately owned reserve and thesite of the Joseph W. Jones Ecological ResearchCenter in Baker County, Georgia, USA (Fig.1). Ichauway is dominated by second growthLongleaf Pine forest and has been intensivelymanaged with prescribed fire since the early1900s. The 11,769-ha property also containsother pines (e.g., Slash Pine, Pinus elliottii,Loblolly Pine, P. taeda, and Short-leaf Pine, P.echinata), hardwoods (typically oaks, Quercusspp.), wildlife food plots for the NorthernBobwhite (Colinus virginianus) and othergame species (Boring 2001), and numeroussemi-permanent to extremely ephemeral wet-lands (Drew et al. 1998; Kirkman et al. 1998;Fig. 1). Two large streams, Flint River andIchawaynochaway Creek, and several pavedhighways intersect the property. Center-pivotagriculture is the primary land use surroundingthe reserve.

Radio-telemetry.—We collected Eastern BoxTurtles opportunistically throughout Ichauwaybetween August 2003 and March 2004. For eachturtle, we measured body mass and straight-linecarapace length (CL) and determined sex basedon the degree of plastron concavity and eye color(Nichols 1939). We individually marked turtlesby notching the marginal scutes with a triangu-lar file (Cagle 1939). We outfitted 23 adult tur-tles (> 100 mm CL) with SM-1H radio trans-mitters weighing 26 g (AVM Instrument Co.Ltd., Livermore, California, USA). Transmitterswere affixed to the costal scutes of the carapacewith epoxy putty (Oatey Supply Chain Services,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 Ichauwayreserve, Baker County, Georgia, USA.

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transmitter and fixative averaged < 40 g (approx-imately 10% of the body mass of the turtle).

We released radio-tagged turtles at the pointof capture and located each turtle at least onceper week during the active season (March-November) and once every two weeks duringthe inactive season (December-March) betweenSeptember 2003 and October 2004. We trackedturtles using an R1000 radio-telemetry receiver(Communications Specialists Inc., Orange,California, USA) and a folding Yagi 6-elementantenna (Wildlife Materials Inc., Murphysboro,Illinois, USA). We radio-tracked turtles untilsignals were lost or the study ended (October2004). We recorded turtle locations using eithera Trimble GeoExplorer 3 (accurate to within 10m) or a TrimbleARC1 (accurate to within 1 m;Trimble Navigation Systems Ltd., Sunnyvale,California, USA). We removed transmitters atthe completion of the study.

Data analysis.—We performed statistical anal-yses with SAS 9.3 software (SAS Institute Inc.,Cary, North Carolina, USA). We set a signifi-cance level of α = 0.05. We performed spatialanalyses with ArcMap 9.3.1 software (ESRI, Red-lands, California, USA). We used an existing ge-ographic information system delineating habitattypes for Ichauway, produced by screen digitiz-ing and ground-truthing 2006 color infrared dig-ital aerial photography at 1-ft pixel resolution.Habitat types were pine forest, hardwood forest,pine-hardwood forest, agricultural, pine planta-tion, aquatic, shrub/scrub, and developed. Pineforest referred to mixed-age stands with > 90%Longleaf Pine (dominant across 90% of this habi-tat type), Slash Pine, Loblolly Pine, or ShortleafPine. Hardwood forest contained > 90% hard-woods including Red Oak (Q. falcata), Live Oak(Q. virginiana), Laurel Oak (Q. laurifolia), orWater Oak (Q. nigra). Pine-hardwood forest wascomposed of 20–80% pines or hardwoods. Agri-cultural land comprised wildlife food plots andother small croplands. Pine plantation consistedof newly planted to mature even-aged stands of

Longleaf Pine or Slash Pine. Aquatic areas wereforested and non-forested isolated wetlands, theIchawaynochaway Creek, and the Flint River.Shrub/scrub included abandoned clearcuts andagricultural fields, failed pine plantations, andhardwood scrub. Developed areas were roads,houses and sheds.

For each turtle, we calculated two estimatesof home range size, 95% minimum convex poly-gon (MCP; Mohr 1947) and 95% kernel den-sity estimate (KDE; Worton 1989), and one es-timate of core area size (50% KDE) using theHome Range Tools extension (HRT; Rodgerset al. 2007) in ArcMap. We selected smoothingfactors (h) that generated 95% KDEs similar inarea to the 95% MCPs (Row and Blouin-Demers2006). We used linear regression (PROC REG)to determine whether home range size estimateswere related to length of the tracking period(days) or number of locations.

We defined the study area as a 2,046-ha areaencompassed by 500-m circular buffers aroundeach MCP. When buffers of individual turtlesoverlapped, we used the Dissolve Tool to cre-ate a single, continuous polygon (Johnson l980;Fig. 1). A small portion of the study area fell out-side of Ichauway in an area for which we had nohabitat data. However, none of the turtles movedoutside of Ichauway, indicating that the compo-sition of the study area with habitat data wasrepresentative of habitat available to the turtles.

We used compositional analysis, a test of pro-portional habitat use relative to availability, toassess habitat selection by turtles, both with andwithout sex-specific comparisons (Aebischer etal. 1993). To assess habitat selection at the lo-cal scale, we compared habitat composition atunique turtle locations (i.e., telemetry locationsin which the turtle had moved > 5 m from itspreceding location) to habitat composition of thehome ranges (95% KDE and 95% MCP), andhabitat composition of the core areas (50% KDE)to that of the home ranges (95% KDE). To as-sess habitat selection at the landscape scale, wecompared both habitat composition of the home

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ranges (95% KDE and 95% MCP) and of the coreareas (50% KDE) to that of the study area (John-son 1980). We performed multivariate analysesof variance (MANOVA) of log-ratio transformeddata (PROC GLM). When a MANOVA indicatedselection, we identified the habitat types selectedby turtles with a ranking matrix of t-tests (PROCMEANS).

We assessed groundcover preferences with use-availability analyses (Neu et al. 1974). Here, wedefine use as the groundcover type at unique tur-tle locations, i.e., whenever a turtle moved >5 m from its preceding location, we recordedthe groundcover type on which it was presentlysitting. Groundcover types were bare ground(no vegetation), litter (leaves and pine nee-dles), grasses (predominantly Wiregrass, Aris-tida stricta, and Broomsedge, Andropogon spp.)and forbs (predominantly oak seedlings, Quer-cus spp., Silkgrass, Pityopsis graminifolia, black-berry, Rubus spp., and Poison Oak, Toxicoden-dron radicans). For analyses, data were summa-rized as percentage of all unique locations withineach of the different groundcover types. We esti-mated groundcover availability with an existingdata set for Ichauway that included percentagesof groundcover types within 442 randomly placed1-m2 quadrats (Miller 2008). We used chi-squarestatistics (PROC FREQ) to test the hypothesisthat groundcover types were used in proportionto their availability. If the null hypothesis wasrejected (α < 0.05), we employed confidence in-tervals following methods in Neu et al. (1974) todetermine which groundcover types were used inproportions greater or less than availability.

Results

We radio-tracked 19 turtles (seven female, ninemale, three of undetermined sex) for at least 4mo of the active season. We excluded four in-dividuals that were lost within 4 mo of track-ing from analyses. We tracked individuals from35–90 times over 174–401 d (x̄ = 326 d; Ap-pendix 1). MCPs were 0.33–54.37 ha and av-

eraged 10.33 ± 3.33 ha. The average MCP formales (11.70 ± 5.75 ha) was larger than thatfor females (8.33 ± 5.68 ha). KDE home rangeswere similar in area to MCPs, as we designed.Core areas were 0.09–10.44 ha and averaged 2.08± 0.62 ha. The average core area for males (2.33± 1.10 ha) was also larger than that for females(1.60 ± 0.99 ha; Appendix 1). Home range sizeestimates were not significantly related to lengthof time tracked (r2 = 0.021, P = 0.554) or numberof tracking events (r2 = 0.139, P = 0.116). Therelatively high degree of variation in MCPs wasdue to two individual turtles with exceptionallylarge home ranges (male 1135, MCP = 54.37ha and female 1142, MCP 42.03 ha, Appendix1). For comparison, the average MCP excludingthese two individuals was 5.87 ± 1.36 ha.

Habitat use did not differ between female andmale turtles (F7,12 = 0.270, P = 0.950), so wereport results of models without a sex compo-nent. At the landscape scale, habitat compositionof the study area differed from that of core ar-eas (F7,12 = 9.94, P < 0.001) and home ranges(KDE: F7,12 = 6.99, P = 0.002; MCP: F7,12 =

10.48, P < 0.001). Specifically, the proportionof mixed pine-hardwood and hardwood forestin home ranges and core areas was greater thanexpected based on availability, whereas the pro-portions of agricultural land, aquatic habitats, de-veloped areas, pine forests, pine plantations, andscrub-shrub habitat in home ranges were less thanexpected (Fig. 2). At the local scale, habitat com-position of home ranges did not differ from that ofcore areas (KDE: F7,12 = 0.96, P = 0.498) or tur-tle locations (MCP: F7,12 = 1.08, P = 0.431; KDE:F7,12 = 2.10, P = 0.123). Eastern box turtle useof groundcover differed from random (χ2 = 32.9,df = 3, P < 0.001); turtles preferred groundcovercomprised of forbs and avoided bare ground andlitter (Fig. 3).

Discussion

To our knowledge, this is the first account ofhome range size and habitat selection of Eastern

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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 similarto MCPs and are not shown.

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

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Box Turtles in a Longleaf Pine landscape matrix.Longleaf Pine-dominated forests at Ichauway andthroughout the Southeastern Coastal Plain aremanaged with frequent prescribed fire. Studies inother regions have reported injuries and mortalityof box turtles from fire (Babbitt and Babbitt 1951;Platt et al. 2010). Our findings indicate, however,that despite risk to individuals from occasionalfires, fire-maintained forests may offer importantbenefits to Eastern Box Turtle populations in thesoutheastern United States.

Home range sizes of Eastern Box Turtles onthe Ichauway reserve were highly variable ashas been reported for the species in other habi-tats (Dodd 2001). Previously reported averageMCPs in the southeastern US are 1.88 ha (Ten-nessee: Donaldson and Echternacht 2005), 2.68ha (North Carolina: Kapfer et al. 2013), and 6.45ha (North Carolina: Hester et al. 2008). Averagehome range size in our study was comparativelyhigh (10.33 ha) owing to one male and one fe-male with home ranges > 40 ha. For the male(#1135; Appendix 1), tracking coincided withtwo long movements across and returning acrossa two-lane highway in April (1,913 m over 8 d inone direction and 1,757 m over 2 d on the return).The female (#1142) moved at least 553 m over3 d in July (184.4 m/d). Box turtles in Georgiamate from April to June and nest in late spring orsummer (Cash and Gibbons 2008); thus, it is pos-sible that these extended movements over a shortperiod of time were related to mate searching andnesting (Stickel 1950; Iglay et al. 2007). Exclud-ing these two individuals, our estimate of MCPhome range size was 5.87 ha, which is closer tothat reported in surveys elsewhere in the south-eastern US (Donaldson and Echternacht 2005,Hester et al. 2008, Kapfer et al. 2013). Addi-tional long-term research is needed to more fullypredict the spatial requirements of the speciesat the regional level and in response to land usechange from urban and agricultural development.However, our home range findings highlight theimportance in conservation planning of protect-ing wildlife corridors and contiguous tracts of

suitable habitat to ensure that large home rangescan be accommodated when they are essential forsurvival and reproduction.

Our compositional analysis revealed habitatselection at the landscape scale but not withinhome ranges, underscoring the value of a two-stage approach to analyses of habitat use (Ae-bischer et al. 2003). Specifically, the turtles thatwe tracked favored pine-hardwood forest andhardwood forest in our study area. On Ichauway,pine-hardwood forest is found on a wide soilmoisture gradient from upland sand ridges towetland edges but is generally characterized asa mesic habitat with a significant componentof Longleaf Pine (Kirkman et al. 2001). Hard-wood forests on Ichauway generally occur inproximity to streams (Fig. 1) on more mesicsoils (Jacqmain et al. 1999; Goebel et al. 2001).The association between Eastern Box Turtles andmesic forests is well supported by previous work(reviewed by Dodd 2001) and is linked to thespecies’ thermal and hydric requirements (Rea-gan 1974; Rossell et al. 2006). Mesic forests typ-ically have dense tree canopies. These canopiescan moderate temperatures and retain understoryhumidity but still permit intermittent sunlight pen-etration and basking opportunities (Dodd 2001).On Ichauway and throughout the SoutheasternCoastal Plain, frequent fire leads to similarities infloral attributes between pine-hardwood forestsand Longleaf Pine savanna. Similarities includediverse native groundcover (Lemon 1949; Walkerand Peet 1984; Kirkman et al. 2001) and highfaunal diversity (Guyer and Bailey 1993), withthe oak component adding canopy cover, verti-cal structure, seedlings, and coarse woody debris.Although the ecological role of oaks in LongleafPine ecosystems is heavily debated, Hiers et al.(2014) contend that increased canopy closure bypyrophytic oaks (i.e., oak species such as Q. fal-cata that are associated with fire-prone uplands)typically does not significantly affect understorydiversity at the stand scale. They also suggestthat presence of pyrophytic oaks may facilitateincreased arthropod diversity and biomass, thus

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potentially increasing habitat quality (e.g., for-age) for some species, including the Eastern BoxTurtle. At a fine scale, turtles in our study pre-ferred patches of forbs and avoided bare ground,litter, and grass substrates. Additional researchis needed to determine whether this groundcoverprovides more suitable foraging and thermoregu-latory environments than other available ground-cover types. If this is the case, it might explainour observation that turtles avoided pine planta-tions and agricultural areas, which lack ground-cover vegetation.

Our results contribute new information to thebody of literature on the habitat requirementsof this species in the southeastern United Statesand shed light on the value of ecological aware-ness in regional forest management. Specifically,although removal of hardwoods is typically amajor element of Longleaf Pine forest manage-ment, our study supports the view that a modestupland-associated oak component can increasethe ecological value of Longleaf Pine forestswhen combined with prescribed fire (Hiers etal. 2014). Knowledge of home range size andhabitat selection of Eastern Box Turtles is criticalfor establishing habitat conservation and restora-tion priorities, especially considering continuingloss of forested habitats to urban and agricul-tural development throughout the range of thisspecies and growing concern for infectious dis-eases (Feldman et al. 2006; Johnson et al. 2008).Given the dominance on the southeastern land-scape of large-scale agriculture and urban/ subur-ban development, we encourage future researchon the habitat preferences of Eastern Box Turtlesin non-forested sites.

Acknowledgements.—This study was partiallyfunded by the University of Florida and theJoseph W. Jones Ecological Research Center.Work was conducted under Georgia Departmentof Natural Resources Scientific Collecting Per-mits 29-WMB-03-142 and 29-WMB-04-84. Wethank Dick Franz and George Tanner from theUniversity of Florida for guidance throughout thefield study. We thank Jean Brock and Jen Howze

for assistance with GIS. Mike Conner and JenHowze also assisted with data analyses and re-viewed an earlier draft of this manuscript. Wealso thank Jones Center staff for collecting turtlesfor this study. This paper was written in memoryof Dr. Larry Galblum.

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

Table

A1.

Body

sizeatcapture

(CL

=carapace

length),numberoftim

esa

turtlew

asradio-tracked,num

berofuniquetracking

locations,lengthof

trackingperiod,and

home

rangesize

(inha)for19

radio-trackedTerrapene

carolinacarolina

usingm

inimum

convexpolygons

(MC

P)andkernel

densityestim

ates(K

DE

),BakerC

ounty,Georgia,U

SA,2003–2004.

F=

female;M

=m

ale;U=

undetermined

sex.

TurtleID

CL

Mass

Telemetry

No.

Unique

Days

95%95%

50%K

DE

(gender)(m

m)

(g)locations

Locations

TrackedM

CP

KD

E(core

area)1132

(F)122

36878

41397

0.870.87

0.221133

(F)144

57071

37365

3.343.37

0.651139

(F)123

36077

41330

7.307.13

1.781142

(F)140

61069

35350

42.0241.62

7.411147

(F)140

56060

22340

3.013.14

0.711151

(F)122

39035

27174

0.330.36

0.093000

(F)130

49176

26322

1.431.51

0.371124

(M)

127329

9050

4012.12

2.000.52

1131(M

)119

30385

45391

5.385.36

1.241135

(M)

127385

4618

24054.37

53.9910.44

1136(M

)132

35971

32371

3.523.69

0.801138

(M)

132415

6329

33620.48

20.753.81

1143(M

)132

45068

33350

13.2613.44

3.131144

(M)

132428

7435

3351.47

1.490.25

1146(M

)109

29670

46339

3.113.05

0.491150

(M)

127390

3526

1741.54

1.560.28

1125(U

)124

37565

33257

11.3811.56

2.371137

(U)

115330

7338

3719.04

8.851.78

1141(U

)107

31777

32343

12.2512.04

3.11

Means

(SE)

Females

132478

8.338.28

1.60(4)

(40)(5.68)

(5.62)(0.99)

Males

126373

11.7011.70

(5.72)2.33

(3)(18)

(5.75)(5.72)

(1.10)

Overall

127407

34326

10.3310.30

2.08(2)

(21)(2)

(15)(3.33)

(3.30)(0.62)

110

Herpetological Conservation and Biology

Sasha E. Greenspan received a B.A. in environmentalscience and anthropology from Wesleyan University andan M.S. in ecology and environmental science from theUniversity of Maine. She is currently working toward aPh.D. in tropical ecology at James Cook University inQueensland, Australia. Her research and career interestsinclude amphibian and reptile ecology and conservation,tropical and disease ecology, One Health, and biologi-cal field station operations. (Photographed by StephenHarris).

Erin P. Condon received her undergraduate degree inEcology and Evolutionary Biology from Princeton Uni-versity and attended graduate school in the Department ofWildlife Ecology and Conservation at the University ofFlorida from 2002–2004. Erin received J.D. and LL.M.degrees from the Frederic G. Levin College of Law andis the former Executive Director of Florida Defendersof the Environment in Gainesville, Florida, USA. (Pho-tographed by Sharon Rush).

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 ofthe effects of mammalian predators on Gopher Tortoises(Gopherus polyphemus), upland snake ecology, and habi-tat predictors of pond-breeding amphibians. Her researchinterests also include the effects of habitat restoration andmanagement on amphibian and reptiles. (Photographedby Miguel Pedrono).

111