Translocated and Resident Eastern Box Turtles (Terrapene c.
carolina) in New York: Movement Patterns and Habitat Use
Megan C. Henriquez, Suzanne K. Macey, Erin E. Baker, Lisa B. Kelly, Rachel L. Betts, Michael J. Rubbo, and J. Alan Clark
Northeastern Naturalist, Volume 24, Issue 3 (2017): 249–266
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2017 NORTHEASTERN NATURALIST 24(3):249–266
Translocated and Resident Eastern Box Turtles (Terrapene c.
carolina) in New York: Movement Patterns and Habitat Use
Megan C. Henriquez1,2, Suzanne K. Macey1,3, Erin E. Baker4,5, Lisa B. Kelly4,
Rachel L. Betts1,6, Michael J. Rubbo4,7, and J. Alan Clark1,*
Abstract - Translocation of animals to new habitats is a common conservation management
strategy but is of uncertain effectiveness. Terrapene c. carolina (Eastern Box Turtle) are
often the subject of translocation efforts. To understand the effectiveness of this strategy,
we radio-tracked 19 translocated and 7 resident Eastern Box Turtles to assess movement
patterns and habitat use, including hibernacula selection. Using data collected over 4 years
from a nature reserve in New York, we compared home range, maximum distance traveled,
and total distance traveled for both translocated and resident turtles. We found no difference
between translocated or resident turtles or between sexes for any of these measures. These
results suggest that translocated turtles at this site adapted well to their new habitat.
Introduction
Many turtle species in North America need to be actively managed because of
increased urbanization (Dodd and Seigel 1991, Ferronato et al. 2015, Gibbs and
Shriver 2002). Urbanization not only decreases the amount of suitable habitat available
for these species, but it also fragments existing habitat (Belzer and Steisslinger
1999) and could lead to inhibition of turtle population growth (Ferronato et al.
2015, Gibbs and Shriver 2002, Steen and Gibbs 2004). Additionally, turtles in
urban and suburban areas are threatened by collisions with vehicles, mowing activity,
and people removing them from the wild either out of a concern for the turtles’
safety or to keep them as pets (Belzer and Steisslinger 1999, Budischak et al. 2006,
Greenspan et al. 2015, Hester et al. 2008). Because turtles have delayed sexual
maturity (Congdon and Gibbons 1990), the loss of individual adult turtles by these
threats may decrease overall local population viability (Belzer and Steisslinger
1999, Budischak et al. 2006, Hall et al. 1999, Kipp 2003).
To mitigate some of urbanization’s negative effects on turtle populations, researchers
and conservation managers often use techniques known as “RRT”, i.e.,
the “repatriation, relocation, and translocation” of individuals, as a conservation
management strategy (Dodd and Seigel 1991, Ferronato et al. 2015, Reinert 1991,
Sosa and Perry 2015). The use of each of these terms by researchers and conservation
managers varies throughout the literature. The International Union for the
1Department of Biological Sciences, Fordham University, Bronx, NY 10458. 2Current address
- Department of Anthropology, The Graduate Center, City University of New York, New
York, NY 10016. 3Center for Biodiversity and Conservation, American Museum of Natural
History, New York, NY 10024. 4Teatown Lake Reservation, Ossining, NY 10562. 5Ramapo
Ridge Middle School, Mahwah, NJ 07430. 6Centre College, Danville, KY 40422. 7Woodcock
Nature Center, Wilton, CT 06897. *Corresponding author - jaclark@fordham.edu.
Manuscript Editor: Peter K. Ducey
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Conservation of Nature defines the term “conservation translocation” as the act of
moving organisms from one area and releasing them in another for conservation
purposes (IUCN SSC 2013). For this study, we use the term “translocation”.
To better understand the effectiveness of translocation efforts, researchers and
conservation managers often track the movement patterns and survivorship of
translocated individuals to investigate the relative success of translocation at an
individual level (Ashton and Burke 2007, Cook 2004, Sosa and Perry 2015). Previous
studies suggest that translocated individuals may have more difficulty acquiring
resources, and, consequently, may have larger home ranges and travel greater distances
than resident turtles in the same area (Attum and Cutshall 2015, Rittenhouse
et al. 2007). The study of movement patterns and home ranges for translocated
turtles can provide researchers with important information not only about dispersal
and survival but also about how translocated individuals use their new habitat and
the quality of that habitat (Greenspan et al. 2015, Kapfer et al. 2013, Stickel 1950).
Terrapene spp. (box turtles) are commonly managed through RRT. In most
published box turtle translocation studies, individuals are drawn from known
populations and released into new habitats (introduction) or habitats supporting an
existing population (reinforcement) (e.g., Cook 2004, Farnsworth and Seigel 2013,
Samuelson 2012). Our reinforcement translocation study of Terrapene c. carolina
(L.) (Eastern Box Turtle), listed as “vulnerable” by the IUCN (van Dijk 2011) and
of “special concern” by New York State (Breisch and Behler 2002, NYSDEC 2017),
is distinctive as it evaluated the effects of translocation on individuals for which
little, if any, information was available on their provenance.
We tracked turtles’ movement patterns and habitat use to better understand
the potential for translocation as a conservation tool for nature reserves and wildlife
rehabilitation centers receiving Eastern Box Turtles whose provenance was
unknown—a common situation for this species. Because translocated turtles are
unlikely to be familiar with their new habitat, we predicted that translocated turtles
would have larger home ranges and travel farther than resident turtles. In addition,
we explored whether translocated turtles spent more time in habitat types different
than those used by resident turtles. We also predicted that differences in movement
patterns between translocated turtles and resident turtles might decrease over time
as translocated turtles gained familiarity with their new habitat. Finally, based on
results of previous studies (e.g., Cook 2004, Kapfer et al. 2013, Stickel 1989), we
did not expect to find a difference in movement patterns and habitat use between
males and females.
Field-site Description
This study was conducted at a private nature reserve in suburban Westchester
County, NY, ~55 km north of New York City. The site was comprised of 338 ha
of mixed habitat, including lakes, deciduous Quercus (oak)–Carya (hickory) forests,
hardwood swamps, and successional shrubland (Fig. 1). The successional
shrubland portion of the site consisted of an open area located under 2 sets of power
lines. The vegetation under the power lines was actively maintained and mowed or
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cut down bi-annually, especially at the bases of the main power line stanchions. The
area surrounding the reserve consisted mostly of private property and residences,
which included manicured lawns, continuation of the oak–hickory forest, and
orchards. For this study, any habitat not on the reserve’s property was considered
“off-property” and not further categorized into specific habitat ty pes.
Methods
Field methods
Resident turtles were found on the reserve prior to the beginning of the study.
Whether resident turtles hatched from nests at the reserve, migrated naturally to
the reserve, or were translocated to the reserve by local citizens at an earlier time
was unknown, and, therefore, we labeled them as resident rather than native. The
nature center at our study site in New York State regularly receives Eastern Box
Turtles from concerned citizens who remove turtles from roads or private property,
from people who no longer want them as pets, and from the New York State Department
of Environmental Conservation (NYSDEC), which confiscates turtles from
the pet trade (box turtles cannot be kept as pets in New York State, pursuant to NY
Environmental Conservation Law §§11-0512). Upon receipt of these turtles by the
Figure 1. Map of study site including habitat types used by Eastern Box Turtles and a subset
of home ranges from one year (selected 2013 individuals, including, but not distinguishing
between, males and females or translocated and resident turtles). White areas represent
“off-property” areas with no habitat data. Shrubland was found only in the power line
rights-of-way.
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nature center, every effort was made to obtain information about the individual's
background and to return it to its habitat of origin. If, however, a turtle’s provenance
could not be determined, we considered it for inclusion in this study. Translocated
turtles were released at different times throughout the 4 years of this study in small
groups (5–7 individuals) at a single designated release point within successional
shrubland habitat on the reserve.
We marked all study turtles, both translocated and resident, using a triangular
metal file to notch the marginal scutes with unique notch codes (modified from
Cagle 1939) and attached small radio transmitters (ATS, R1850, Isanti, MN) to the
back-right marginal and costal scutes using waterproof epoxy (Oatey, Cleveland,
OH) which was then rubbed with soil to obtain a more natural color and smell. The
total weight of the transmitter and epoxy was less than 5% of the turtles’ weight. The transmitters’
antennae were not glued down, were 12.5 cm long, and came off the shell at
a 15-degree angle facing the tail. Transmitter attachment was completed within 30
minutes. For resident turtles, we attached transmitters in the field and released the
individuals at their capture location. For translocated turtles, we attached transmitters
at the reserve’s on-site wildlife center prior to their release onto the reserve. All
capture and study procedures were conducted under a NYSDEC Scientific License
to Collect or Possess.
Tracking
To evaluate Eastern Box Turtle movement patterns and habitat use, we tracked
26 turtles for up to 4 years (2010–2013). Nineteen turtles were translocated (9
males and 10 females), and 7 turtles were residents (3 males and 4 females). We
tracked turtles 2–3 times per week during their yearly active period, which we
defined as the time they emerged from their hibernacula (~April), to the time they
buried themselves underground for the winter (~November). We used a handheld
R-1000 receiver and an RA-150 Yagi antenna (Communication Specialist, Inc.,
Beacon, NY) to locate individuals and a handheld GPS unit (Garmin GPS map60,
accuracy less than 15 m) to record their location. If we were unable to track turtles multiple
times per week, we checked their radio signal weekly to ensure that the transmitter
was still working and the individual was still within range.
During tracking events, any turtle that showed signs of disease or injury was
brought to an on-site health facility and rehabilitated until deemed healthy by a certified
wildlife rehabilitator. After rehabilitation, turtles were released at their last point
of capture or another location within their established home range. When a turtle was
found in an area considered unsafe (e.g., a road), we moved the turtle to its nearest
previous tracking location. We tracked each turtle for the entire winter, albeit less
frequently once an individual went below the soil surface, and considered the last location
recorded for that calendar year to be the hibernaculum point for that year.
Movement patterns
Because of seasonal variation in Eastern Box Turtle activity (Dolbeer 1969, Stickel
1950) and the variation in our yearly tracking of study individuals, we standardized
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the tracking data by using the same 20-week period (21 May–22 October) in all years
of this study for the movement-pattern analysis. We define “tracking season” as this
standardized 20-week period. Any turtle that was not located for 3 or more consecutive
weeks at any time during the tracking season was excluded from all analyses for
that year. We examined home range, total distance traveled, and maximum distance
traveled for each tracking season for each individual and also averaged these measurements
across all tracking seasons an individual was observed.
We used global positioning system (GPS) coordinates to map and calculate
each turtle’s home range using the Minimum Convex Polygon (MCP) function in
ArcGIS 10.0 (ESRI 2011). Although some studies used fixed kernel density and
harmonic means to estimate home range (e.g., Cook 2004, Kapfer et al. 2013,
Refsnider et al. 2012), we used the MCP method because it is more commonly
used in box turtle movement studies and so allowed us to more easily compare our
results to these studies (see Appendix 2). To calculate the total distance traveled,
we summed the distance traveled between each consecutive tracking event. To calculate
the maximum distance traveled, we measured the greatest distance between
any 2 (not necessarily consecutive) points in each turtle’s home range.
We visually inspected and tested residual plots of the tracking season’s movement
data for normality using the Shapiro-Wilkes test (shapiro.test function) in
the stats package for R (R Code Team 2014). Because home range, total distance
traveled, and maximum distance traveled were non-normal and residual variance
increased with the mean, we log transformed movement data to meet the assumptions
of normality. The final comparisons of movements were made using the linear
mixed effect analysis (lmer function) in the lme4 package (Bates et al. 2014).
In the linear mixed model, we set sex (i.e., male and female) and status (i.e.,
translocated or resident) as fixed effects. We set individual, calendar year, and
number of tracking points as random effects. We compared each candidate model
(i.e., sex and status interaction, sex and status, sex alone, status alone) to the null
using a likelihood ratio test (anova function, stats package), and we compared candidate
models to each other by creating AICc tables using the aictab function in the
AICcmodavg package. For models significantly different than the null model, but
not distinguishable from other candidate models, we performed model averaging
to assess the effect of the parameters given model uncertainty (modavg function,
AICcmodavg package).
To detect trends in movement patterns over time for translocated turtles, we
analyzed movement patterns considering the number of years (post-release) the
turtle was present at the site (site year). Of the 19 translocated turtles tracked, 15
had adequate tracking data over multiple tracking seasons to be considered for this
analysis, and we ran a separate set of linear mixed models on the movement data of
these individuals. In these models, all of the above-mentioned random effects were
maintained to account for differences in individuals, calendar year, and number
of seasonal tracking points collected. The candidate models included sex and site
year interaction, sex and site year, sex alone, and site year alone; a comparison of
models was performed with the same approach as described above.
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Habitat use
To assess relative habitat use for all study turtles, we used ArcGIS to superimpose
GPS tracking locations onto a data layer of the habitat types for the reserve
(i.e., oak–hickory forest, wet meadow, vernal pool, upland meadow, successional
shrubland, hardwood swamp, shallow lake, and rocky slope). The habitat layer was
constructed using aerial photography to delineate the different habitat types and
was verified in the field.
We conducted a compositional analysis to study habitat use at the study
site, with the compana function using the parametric test in the adehabitatHS
package for R (Calenge 2006). This analysis takes into consideration both
the frequency of habitat use for each individual and the amount of habitat available
within the reserve (Aarts et al. 2008, Aebischer et al. 1993). Any turtle
position not on the reserve was considered “off-property” and not included in
the compositional analysis as the description and proportion of off-property
habitat was unknown.
Compositional analyses cannot be used to compare differences between groups
of individuals (e.g., males vs. females; Aebischer et al.1993). Therefore, for the
2 habitat types used most (based on the results of the compositional analysis),
we performed generalized linear mixed models (glmer function; binomial family
with logit link function; lme4 pacakage) to assess differences of habitat use among
groups. The candidate models included sex and status interaction, sex and status,
sex alone, and status alone; comparison of models was performed with the same
approach as described above for the movement analysis.
Overwintering behavior
Box turtles often dig shallow cavities into the ground or use natural pits for hibernating
during the winter (i.e., hibernacula) when food is scarce and temperatures
are prohibitive to more active metabolic processes (Ultsch 2006). When turtles are
translocated into new environments, finding appropriate hibernaculum sites may
be challenging, as these turtles are not familiar with local habitat conditions (but
see Cook 2004, Farnsworth and Seigel 2013). In New York, temperatures regularly
drop below freezing, and securing a good hibernaculum is critical to a turtle’s survival
(Carpenter 1957, Claussen et al. 1991). Therefore, we also tracked turtles’
overwintering behavior, including their selection of hibernacula.
To evaluate overwintering behavior, we recorded the habitat type in which each
hibernaculum site was located based on the habitat layer used in the habitat-use
analysis. We also noted the frequency with which each hibernaculum habitat type
was used, year to year, by each individual turtle. To assess hibernaculum fidelity, we
defined fidelity to occur when a turtle buried itself less than 15 m from the previous year’s
hibernaculum. The choice to use a measurement of less than 15 m was based on other studies
(e.g., Refsnider et al. 2012, Stickel 1989) as well as our GPS unit’s accuracy,
which was limited to less than 15 m.
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Results
Tracking
The number of individual turtles whose tracking data were used for the study
varied between years (2010 and 2011: n = 12; 2012: n = 20; 2013: n = 18) due to
tracking inconsistency within the 20-week tracking season, transmitter failure/
availability, or mortality. Only 3 turtles (FRes1, FTrans4, and MRes2) in this study
had data for all 4 tracking seasons (Appendix 1). However, an additional 3 turtles
(FRes2, MRes3, MTrans4) were tracked over the entire 4-year study period, but
only 3 tracking seasons within the 4-year period were included because of lack of
sufficient data for 1 of the tracking seasons (Appendix 1). All measures and results
are given ± S.D. Most individuals were tracked for 2 or more calendar years (mean
= 2.4 ± 1.0 years). We collected 1721 tracking points during this study, and the
number of points for an individual turtle during a tracking season varied from 16 to
61 points (mean = 28 ± 12.1 points) per tracking season across years.
Movement patterns
During the 20-week tracking season, individual home ranges varied from 0.03
to 79.8 ha (mean = 8.4 ± 15.0 ha), while individual averages across tracking seasons
ranged from 0.07 to 50.6 ha (mean = 8.2 ± 12.0 ha) (see 2013 home ranges in Fig. 1;
all home ranges given in Appendix 1). Total distance traveled, or the sum of the
distances between all tracking points collected per individual in 1 tracking season,
varied from 280 to 13860 m (mean = 2247.6 ± 2529.8 m). Average total distances,
or the average of all the tracking season total distances collected for each individual,
ranged from 332 to 6787 m (mean = 2253.6 ± 1833.2 m; Appendix 1). Maximum
distance, or the longest distance between any 2 points per individual in one tracking
season, varied from 28 to1451 m (mean = 442.5 ± 342.5 m). Average maximum
distances, or the average of all tracking season maximum distances collected per
individual, ranged from 42 to 1441 m (mean = 590.3 ± 565.4 m) (Appendix 1). Based
on the results of the linear mixed model comparisons, none of the candidate models
in any movement category were better performing than the null model (Table 1).
For translocated turtles, the amount of time (site year) after release did not appear
to affect home ranges, although some individuals had large changes in home
ranges from year to year (Table 2). For example, MTrans9 had an increase of 32.9
ha (284%) between site year 1 and site year 2, while MTrans2 had a 16.3-ha (80%)
decrease in home range between site year 1 and site year 2. MTrans2’s home range
then stabilized for the third year, keeping a ~3.9-ha home range for both the second
and third year after release. This stabilization of home range over time was not
observed in FTrans2, who had an 11.9-ha (75%) decrease in home range between
site year 1 and 2 and then a 7.3-ha (187%) increase between site year 2 and 3. Not
including these 3 individuals with relatively large home-range changes, the average
change (either increasing or decreasing) of home range over 2 consecutive site
years was 1.8 ± 1.9 ha. When comparing candidate models for movement patterns
in translocated turtles, neither sex, site year, nor the interaction between sex and
site year had a significant effect on movement patterns over time (Table 2).
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Table 1. Eastern Box Turtle movement model comparisons based on sex and status as translocated or
resident. Models are listed in order of increasing AICc values. K is the number of estimable parameters
(degrees of freedom), AIC is the Akaike Information Criteria, AICc is the AIC corrected for small
sample size, ΔAICc is the difference in AICc from the “best” performing model, AICc Wt is the relative
weight of each model, and P is the P-value from the likelihood ratio test comparing the candidate
model to the null model (random effects only). No models were considered different than the null.
Model K AIC AICc Δ AICc AICc Wt P
Home range
Status 6 186.8 188.30 0.00 0.43 0.624
Sex 6 186.9 188.41 0.11 0.19 0.718
Sex + status 7 188.6 190.71 2.41 0.13 0.828
Sex : status 8 190.2 192.94 6.85 0.02 0.850
Maximum distance
Sex 6 160.0 161.55 0.00 0.43 0.563
Status 6 160.2 161.72 0.17 0.39 0.687
Sex + status 7 161.9 163.93 2.38 0.13 0.779
Sex : status 8 162.9 165.65 4.09 0.05 0.698
Total distance
Status 6 160.9 162.38 0.00 0.45 0.429
Sex 6 161.4 162.89 0.52 0.34 0.742
Sex + status 7 163.2 164.80 2.42 0.13 0.688
Sex : status 8 163.2 165.89 3.51 0.08 0.512
Table 2. Translocated Eastern Box Turtle movement model comparisons based on sex and site year.
Models are listed in order of increasing AICc values. K is the number of estimable parameters (degrees
of freedom), AIC is the Akaike Information Criteria, AICc is the AIC corrected for small sample size,
ΔAICc is the difference in AICc from the “best” performing model, AICc Wt is the relative weight of
each model, and P is the P-value from the likelihood ratio test comparing the candidate model to the
null model (random effects only). No models were considered different than the null.
Model K AIC AICc Δ AICc AICc Wt P
Home range
Sex 6 122.4 124.64 0.00 0.92 0.936
Site Year 8 125.9 130.04 5.40 0.06 0.928
Sex + Site Year 9 127.9 133.21 8.56 0.01 0.976
Sex : Site Year 12 133.1 143.12 18.47 0.00 0.987
Maximum distance
Sex 6 119.0 121.29 0.00 0.94 0.312
Site Year 8 123.5 127.58 6.29 0.04 0.902
Sex + Site Year 9 124.4 129.73 8.44 0.01 0.808
Sex : Site Year 12 125.9 135.99 14.70 0.00 0.526
Total distance
Sex 6 113.3 115.57 0.00 0.77 0.747
Site Year 8 114.3 118.41 2.85 0.19 0.376
Sex + Site Year 9 116.1 121.42 5.85 0.04 0.512
Sex : Site Year 12 120.4 130.48 14.91 0.00 0.661
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Habitat use
Considering the amount available of each different habitat type, the turtle location
data collected during the tracking season indicated that turtles generally
preferred certain habitats (i.e., successional shrubland and oak–hickory forest)
to others ( λ = 6.9e-16, df = 9, P < 0.001; Table 3). Although the successional
shrubland was found only under the power lines and was less available than the
oak–hickory forest, successional shrubland was the most frequently used habitat by
translocated turtles (Table 4).
When investigating differences in successional shrubland habitat use considering
sex and status, the model with status alone was significantly different than the
null model but not different from the model with both sex and status (Table 5).
When predictor variables were averaged across models, translocated turtles do
not use successional shrubland more than resident turtles (95% unconditional CI
of the model-averaged estimate of status includes zero; Fig. 2), but translocated
turtles do use oak–hickory forest habitat significantly less than resident turtles
(Table 5, Fig. 2).
Table 4. Habitat type and percentage (± SD) use by Eastern Box Turtles organized by status as translocated
or resident and sex (2010–2013). Any habitat types not used by study turtles are not included.
Habitat types are listed by most to least used by all individuals pooled.
Habitat All individuals Resident Translocated Female Male
Shrubland 44.25 ± 0.25 33.44 ± 0.24 48.67 ± 0.25 40.37 ± 0.23 48.96 ± 0.28
Forest 37.20 ± 0.18 43.44 ± 0.18 34.65 ± 0.17 37.89 ± 0.17 37.89 ± 0.19
Off-property 8.89 ± 0.20 6.69 ± 0.12 9.79 ± 0.23 12.12 ± 0.25 12.12 ± 0.10
Swamp 4.75 ± 0.13 13.46 ± 0.22 1.19 ± 0.04 6.79 ± 0.17 2.28 ± 0.06
Rocky slope 3.93 ± 0.11 2.11 ± 0.05 4.68 ± 0.13 2.27 ± 0.05 5.96 ± 0.15
Vernal pool 0.63 ± 0.02 0.66 ± 0.02 0.61 ± 0.02 0.23 ± 0.01 1.11 ± 0.02
Lake 0.29 ± 0.01 0.21 ± 0.21 0.32 ± 0.01 0.34 ± 0.01 0.23 ± 0.01
Wet meadow 0.06 ± 0.00 0.00 ± 0.00 0.08 ± 0.01 0.00 ± 0.00 0.13 ± 0.01
Table 3. Comparisons of habitat use for individual Eastern Box Turtles over all years sampled via compositional
analysis. Table reads left to right, and sign relates to the comparative relationship between
habitat types (i.e., - is used less, + is used more). Triple signs mean statistical difference (P < 0.05) in
the sign direction. Area of habitat and percentage of that habitat type within the study site are listed
under habitat type abbreviations.
Habitat type
SH F SW RS VP L WM
Area (ha) 14.4 243.6 22.5 19.1 1.3 23.4 1.5
% 4.2 71.3 6.6 5.6 0.4 6.8 0.4
Shrubland (SH) +++ +++ +++ +++ +++ +++
Forest (F) +++ +++ +++ +++ +++
Swamp (SW) - - +++ +
Rocky slope (RS) - +++ +
Vernal pool (VP) +++ +++
Lake (L) ---
Wet meadow (WM)
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Table 5. Eastern Box Turtle habitat-use model comparisons based on status as translocated or resident
and sex for shrubland and forest habitat types. Models are listed in order of increasing AICc values. K
is the number of estimable parameters (degrees of freedom), AIC is the Akaike Information Criteria,
AICc is the AIC corrected for small sample size, ΔAICc is the difference in AICc from the “best”
performing model, AICc Wt is the relative weight of each model, P is the P-value from the likelihood
ratio test comparing the candidate model to the null model (random effects only).
Model K AIC AICc Δ AICc AICc Wt P
Shrubland
Status** 5 412.4 413.44 0.00 0.49 0.049*
Sex + status** 6 413.1 414.61 1.17 0.27 0.076
Sex 5 414.9 415.99 2.54 0.14 0.253
Sex : status 7 414.5 416.53 3.09 0.01 0.123
Forest
Status** 5 391.4 392.44 0.00 0.45 0.003*
Sex : status** 7 391.3 393.35 0.91 0.28 0.005*
Sex + status** 6 392.0 393.55 1.10 0.26 0.006*
Sex 5 399.0 400.06 7.61 0.01 0.295
*Models statistically different compared to null.
**Models with less than 2 ΔAICc from the “best” model and therefore model averaging was p erformed.
Figure 2. Comparisons of predictor variables’ model-averaged coefficients with 95% unconditional
confidence intervals (CI) for successional shrubland and oak–hickory forest habitat
use by Eastern Box Turtles. When 95% CI did not include zero (did not cross the dashed
line), the predictor variable was considered to have an effect on habitat use. The direction
of the effect is based on the comparison of translocated turtles to resident turtles for status
and males to females for sex and on the sign of model-averaged coefficient; for example,
translocated turtles use forest less than resident turtles (as represented by the negative
model-averaged coefficient and the CI range not including zero).
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Overwintering behavior
We collected multiple years of hibernacula data for 22 of 26 turtles. Turtles overwintered
in only 2 habitat types: oak–hickory forest and successional shrubland
located under power lines. The oak–hickory forest was the preferred overwintering
habitat, with 17 of 22 (77%) turtles hibernating there for at least 1 winter. Six of 7
(86%) resident turtles and 11 of 15 (73%) translocated turtles hibernated in oak–
hickory forest at least once. Seventeen of 22 (77%) hibernating turtles exhibited
fidelity to habitat type with 15 of those 17 (88%) selecting oak–hickory forest for
each year observed. The average distance between any 2 hibernaculum locations
across all individuals was 124.9 ± 125.7 m. Only 3 of 22 (14%) showed hibernaculum
fidelity (within 15 m of a previous hibernaculum site), including 2 translocated
turtles (FTrans6 and MTrans4) and 1 resident turtle (FRes2).
Discussion
Movement patterns
Home range. Consistent with previous studies of Eastern Box Turtles (Cook
2004, Kapfer et al. 2013, Stickel 1989), we found no difference in home ranges between
males and females. Unlike most previous studies comparing translocated and
resident box turtles (Appendix 2), the home ranges of our translocated turtles were
not larger than the home ranges of our resident turtles. Differences in home range
in other studies may reflect differing analytical approaches, population densities, or
habitat quality at different sites (Appendix 2; Stickel 1950, 1989). When comparing
the results of other studies to ours, we note that our data were based on a 20-week
tracking season that may not have captured all of a turtle’s movements during the
entire active season or be directly comparable to other studies that used different
tracking periods.
In addition to differing tracking periods, the large variability of home ranges for
translocated turtles in our study could reflect differences in the origins of the turtles
themselves, as the provenance of our translocated turtles was unknown. Cook
(2004), however, saw no differences between movement patterns for released pets
versus translocated wild Eastern Box Turtles of known provenance. After translocated
turtles acclimatize to their new environment, home ranges might be expected
to decrease and stabilize over time (Hester et al. 2008); however, we found no
pattern of change in home ranges over time. The large differences in home ranges
between years exhibited by a few turtles may be due simply to individual variation
in home range stability, as seen in Nichols (1939).
Total distance traveled. Only a small number of turtles (both translocated and
resident) in our study showed irregular movement patterns, such as long unidirectional
movement. For example, a resident male showed strong linear movement
(MRes1= 600 m) alternating between 2 smaller sub-home ranges during the course
of each of tracking season. In both our study and Hester et al.’s (2008) study, the
average total distance traveled by translocated and resident turtles did not differ.
However, in both status categories, our turtles traveled ~1 km less than Hester et
al.’s population, which may reflect the shorter time period we used for movement
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analysis within a year (20-weeks), differences in habitat quality, or differences in
resource distribution at each site.
Maximum distance traveled. Translocation projects may be hindered by animals
leaving the study site in their attempts to return to their habitat of origin, a process
known as homing (Berry 1986, Rittenhouse et al. 2007) or because the target site for
the translocation is too small or of too poor quality for the persistence of additional
turtles (Cook 2004, Dodd and Seigel 1991). Translocation studies often attempt to
capture dispersal events by tracking the maximum distance traveled and orientation
of the dispersal (e.g., Cook 2004, Lemaku 1970, Rittenhouse et al. 2007). Based on
the maximum dispersal distance recorded in his study (113–1295 m), Cook (2004)
concluded that a translocation site should be at least 300 ha; our findings on the
average maximum distance traveled for translocated turtles (42–1441 m) support
this recommendation.
Habitat use
Preferred habitat type. Eastern Box Turtles are generally considered a forestdwelling,
terrestrial species (Budischak et al. 2006, Dodd 2001, Williamson 2013).
For example, Williams and Parker (1987) reported resident Eastern Box Turtles in
Indiana strongly favored forest habitat and were rarely seen in power line rightsof-
way. In our study, we also found that resident turtles used forest habitat most
frequently but found that translocated turtles used the successional shrubland habitat
under the power line rights-of-way most frequently.
Our results suggest that translocated individuals may be using the reserve’s habitat
differently than residents, but we acknowledge the reduced use of oak–hickory
forest may be associated with the translocation release point being in the successional
shrubland. Our choice for a release point possibly affected the translocated
turtles’ habitat selection and home-range establishment (Brichieri-Colombi and
Moehrenschlager 2016). Even so, both resident and translocated turtles preferred
to hibernate in the oak–hickory forest which, for translocated turtles, was outside
the habitat type of their release point.
Power lines are ubiquitous in areas affected by urbanization, and our translocated
turtles’ use of habitat beneath power lines raises interesting questions.
While power lines could provide quality habitat for box turtles, the active management
of powerline rights-of-way could also pose a threat. At our site, the
local electric company mowed the areas below power lines bi-annually to avoid
overgrowth of grasses, shrubs, and trees. Although none of our study turtles were
injured or killed by this maintenance, previous studies have shown that mortality
could result from mowing or harvesting grasses (Nazdrowicz et al. 2008).
Consequently, if turtles preferentially use these actively managed areas, they may
experience higher mortality.
Off-property use. No turtles migrated away from the study site, though several
individuals temporarily left the borders of the reserve. Turtles in unprotected offproperty
habitat, such as yards and agriculture fields, may be more vulnerable to
mortality events due to mowing, collisions with vehicles, and collection (Greenspan
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et al. 2015). Although off-property was the third most frequently used habitat in this
study, the unprotected off-property habitat at our site often included high quality
continuations of oak–hickory forest or successional shrubland; as a result, the use
of off-property habitat may not have conferred greater risks to individuals.
Overwintering behavior
Other Eastern Box Turtle studies showed variability in hibernaculum fidelity
(e.g., Claussen et al. 1991, Cook 2004). For example, Stickel (1989) found that
Eastern Box Turtles selected hibernacula sites from 8 to 24 m from previous years’
hibernacula sites. Cook (2004) reported distances between consecutive hibernacula
averaged 97.6 ± 89.9 m, but he described the variability not only between individuals
in a population, but also between years for a single individual. These results are
comparable to our own findings of high levels of individual variability in hibernaculum
selection.
Conclusions
Our study found no differences between home ranges and movement patterns of
translocated and resident turtles. However, the translocated turtles in our study did
not spend as much time in the oak–hickory forest habitat as resident turtles. This
difference could be associated with the release point for our translocated turtles,
which was in successional shrubland. Future translocation projects should carefully
consider release locations.
Translocation sites need to include enough space for relocated individuals to
have sufficient access to food, reproductive opportunities, and other factors critical
to the species’ survival at all life stages (Dodd and Seigel 1991). The size of our
site and the variety of available habitat types may have allowed for translocated
individuals to successfully acquire such resources.
Nature reserves and wildlife rehabilitation centers often receive animals whose
provenance was unknown. Our results suggest that Eastern Box Turtles can be
successfully translocated to new habitats, even if their provenance is unknown. A
limitation of our study is that we were not able to determine whether the resident
turtles in our study were hatched on-site or whether they were previously released
on-site by well-meaning private citizens. Perhaps through rigorous marking programs
or genetic analyses, future studies could know with more certainty the origin
of both resident and translocated turtles.
Only 1 study turtle died during the 4 years of this study: a young translocated
male (MTrans1) displaying evidence of winter exposure and dehydration (Appendix
1). Other studies of translocated box turtles had higher mortality rates
(Farnsworth and Seigel 2013, Hester et al. 2008). Because the reserve had a
wildlife rehabilitator on site, we were able to readily address health issues presented
by either translocated or resident turtles. These healthcare measures might
account for some of the difference between our study and others that had higher
study turtle mortality. Even if consistent health monitoring is not possible for future
translocation studies, we strongly recommend that turtles be quarantined and
tested for disease prior to release, especially considering the increased concern
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about and documentation of disease outbreaks such as Ranaviris in box turtles
(Kimble et al. 2017).
While the definition of a successful RRT project is unclear, many researchers
and managers agree that successful reproduction and incorporation of translocated
individuals into existing populations are some of the primary qualifiers (Burke
1991, Dodd and Seigel 1991, Brichieri-Colombi and Moehrenschlager 2016). In
this study, we did not track reproductive patterns on either the individual or population
level. Future translocation studies would benefit from following individuals
for longer periods of time, while focusing on survivorship, mating patterns, reproduction,
population stability, and the implications translocations have on the health
and genetic structuring of resident populations.
Nature centers and wildlife rehabilitation centers will likely continue to acquire
individual Eastern Box Turtles whose provenance is unknown. The ability to successfully
relocate such individuals may be important for this species’ conservation,
and this study suggests that such translocations have the potential to be a useful
conservation tool.
Acknowledgements
We thank the staff at the nature center at our study site for logistical support and the
nature center’s wildlife center for receiving and rehabilitating turtles, while providing
consistent support with field data collection. We are also grateful to Karina Polanco and
James Herrera for assistance with data analysis. This project was supported by Fordham
University, Fordham University’s Calder Summer Undergraduate Research program, and
the National Science Foundation Research Experience for Undergraduates program.
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Appendix 1. Sex, status, health notes, and average movement measurements over multiple
tracking seasons taken for all Eastern Box Turtles in this study. Turtle ID corresponds to
sex (F = female; M = male), status (Res = resident; Trans = translocated), and individual ID
number. The lower portion of the table is data summarized based on categories of turtles. #
of seasons = number of seasons tracked. In health notes, treated indicates individuals that
were treated for upper respiratory tract infections, and the 1 individual who died likely succumbed
to exposure during the overwintering period. Measurements are given ± SD.
# of Average home Average total Average max.
Turtle ID seasons Health notes range (ha) distance (m) distance (m)
FRes1 4 14.44 ± 10.94 2701 ± 1550 613 ± 259
FRes2 3 1.18 ± 0.98 786 ± 369 215 ± 107
FRes3 1 5.16 1631 374
FRes4 1 0.61 813 173
FTrans1 3 2.90 ± 1.48 1515 ± 188 417 ± 235
FTrans2 3 10.31 ± 5.99 1874 ± 1015 581 ± 236
FTrans3 3 Treated 2012 0.61 ± 0.26 723 ± 539 131 ± 17
FTrans4 4 3.02 ± 1.21 1816 ± 1141 449 ± 238
FTrans5 3 3.05 ± 1.59 1848 ± 538 427 ± 178
FTrans6 3 2.01 + 0.64 2461 ± 1717 426 ± 315
FTrans7 2 3.90 ± 0.79 1523 ± 127 333 ± 92
FTrans8 2 4.42 ± 5.50 1389 ± 1388 293 ± 248
FTrans9 1 2.10 1404 370
FTrans10 1 48.04 4437 1441
MRes1 2 21.10 ± 25.57 6787 ± 7941 742 ± 589
MRes2 4 Treated 2012, 2013 28.78 ± 35.82 6301 ± 5959 831 ± 523
MRes3 3 1.03 ± 0.77 1137 ± 445 156 ± 53
MTrans1 2 Died 2011 0.84 ± 1.15 1041 ± 1077 151 ± 179
MTrans2 3 9.31 ± 9.36 3268 ± 1787 743 ± 51
MTrans3 3 Treated 2012 1.05 ± 0.61 1009 ± 524 223 ± 35
MTrans4 3 Treated 2010 0.57 ± 0.67 504 ± 279 119 ± 67
MTrans5 2 Treated 2013 7.23 ± 4.35 2225 ± 1068 456 ± 167
MTrans6 2 Treated 2013 1.08 ± 0.84 908 ± 408 213 ± 151
MTrans7 1 0.07 332 42
MTrans8 1 50.57 6134 1142
MTrans9 2 28.06 ± 23.29 4026 ± 1278 818 ± 448
Residents (n = 7) 1 treated 10.33 ± 11.29 2880 ± 2590 443 ± 283
Translocated (n = 19) 4 treated, 1 died 9.43 ± 15.42 2026 ± 1488 462 ± 359
Females (n = 14) 1 treated 7.27 ± 12.35 1784 ± 952 446 ± 318
Males (n = 12) 4 treated, 1 died 12.47 ± 16.22 2806 ± 2437 469 ± 368
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Appendix 2. Comparison of home ranges in box turtle (Terrapene sp.) movement studies.
Difference
Average MCP (ha) between
Species Translocated Resident groups? Analytical method Study
Terrapene c. carolina (Eastern Box Turtle) 9.43 10.33 No MCP This study
9.80 - - 95% bivariate normal Cook 2004
4.80 - - 95% harmonic mean Cook 2004
14.71 4.31 Yes MCP Farnsworth and Seigel 2013
18.02 6.45 Yes MCP Hester et al. 2008
- 2.68 - MCP Kapfer et al. 2013
Terrapene c. triunguis (Agassiz) 14.22 7.89 No MCP Samuelson 2012
(Three-Toed Box Turtle)
Terrapene ornate (Ornate Box Turtle) - 0.40 - 50% fixed kernel Refsnider et al. 2012