Effects of Site, Year, and Estimator Choice on Home

Ranges of Bog Turtles ( Glyptemys muhlenbergii) in

Maryland

Authors: Byer, Nathan W., Smith, Scott A., and Seigel, Richard A.
Source: Journal of Herpetology, 51(1) : 68-72
Published By: Society for the Study of Amphibians and Reptiles
URL: https://doi.org/10.1670/15-150

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 Journal of Herpetology, Vol. 51, No. 1, 68–72, 2017
Copyright 2017 Society for the Study of Amphibians and Reptiles

Effects of Site, Year, and Estimator Choice on Home Ranges of Bog Turtles (Glyptemys
muhlenbergii) in Maryland

NATHAN W. BYER,1,2 SCOTT A. SMITH,3 AND RICHARD A. SEIGEL1

1
Department of Biological Sciences, Towson University, Towson, Maryland, USA
3
Wildlife and Heritage Service, Maryland Department of Natural Resources, Wye Mills, Maryland, USA

ABSTRACT.—Bog Turtles (Glyptemys muhlenbergii) are some of the most imperiled turtles in North America, and because of threats
posed by habitat loss and collection for the pet trade, the species currently is listed as threatened under the U.S. Endangered Species Act.
Numerous studies have investigated home ranges and movements of this species, and reported home-range sizes have varied widely.
Typically, these studies were conducted at different sites and used different home-range estimators, and these factors may influence home
ranges. Few studies have assessed the impact of estimator choice on home-range size for Bog Turtles or how habitat changes have affected
their movements. Our study investigated home-range sizes of female Bog Turtles at two sites in Maryland, using multiple home-range
estimators. Female turtles were tracked using radiotelemetry between April and August in 2013 and 2014. Home-range sizes differed
significantly between study sites but did not differ significantly between years. Choice of estimator had a significant effect on home-
range size; 50% kernel and 95% minimum convex polygons were similar, but 95% kernels were significantly larger than 50% kernels or
95% polygons. A study conducted at one of our two study sites over 15 years ago reported smaller home-range sizes than were found in
our study, perhaps indicating that habitat has deteriorated at that site.

Bog Turtles (Glyptemys muhlenbergii; Holman and Fritz, 2001)
are considered to be among the most imperiled North American
turtle species (Chase et al., 1989; Morrow et al., 2001). They are
habitat specialists that use small, spring-fed bogs and fens,
habitats that are often negatively impacted by anthropogenic
habitat modification, proliferation of invasive species, and
natural succession (Chase et al., 1989; Ernst and Lovich, 2009;
Pittman and Dorcas 2009). In addition to habitat loss caused by
these factors, Bog Turtles have been heavily impacted by
collection for the pet trade (Lee and Norden, 1996). Bog Turtle
populations in the northern part of this species’ geographic
range were listed as threatened in 1997 under the U.S.
Endangered Species Act, with southern populations listed as
threatened attributable to similarity of appearance.
Movement patterns and home ranges of this species have
been studied frequently over the past 40 years (e.g., Ernst, 1977;
Chase et al., 1989; Carter et al., 1999; Morrow et al., 2001) and
have provided a great deal of information about the spatial
ecology of these elusive turtles. Home-range sizes for this
species generally are small, with Chase et al. (1989) reporting
harmonic mean home-range sizes of 1.33 ha for males and 1.26
ha for females at a site in Maryland. Home-range sizes also
differ among studies, with Morrow et al. (2001) reporting much
smaller average home ranges of 0.04 ha and 0.07 ha for males
and females, respectively, at two sites in Maryland using the
minimum convex polygon home-range estimator. Although
such differences in home-range sizes could be attributable to the
use of different home-range estimators in each study, Carter et
al. (1999) suggested that differences could also be attributable to
different wetland sizes and, therefore, different availabilities of
microhabitats at each site. This suggests that studies should 1)
use multiple home-range estimators and 2) take into account
differences in wetland sizes between sites. However, few studies
have investigated the impact of estimator choice on home-range
size for Bog Turtles.
2
Corresponding Author. Present address: Department of Forestry
and Wildlife Ecology, University of Wisconsin-Madison, Madison,
Wisconsin, USA; E-mail: nbyer@wisc.edu
DOI: 10.1670/15-150
Our study aimed to use several commonly used home-range
estimators to calculate home-range sizes of Bog Turtles in
Maryland. We investigated two main questions: 1) How do
home-range sizes differ between sites and years for female Bog
Turtles in Maryland? and 2) How does the choice of estimator
affect home-range estimates? Because of funding constraints
posed by a concurrent study of nesting ecology (NWB, unpubl.
data), we used only female Bog Turtles in this study. Therefore,
we did not assess the effect of sex on home-range size, a topic
that has been studied several times (e.g., Carter et al., 1999;
Morrow et al., 2001; Pittman and Dorcas, 2009).
MATERIALS AND METHODS
Study Locations.—We selected study sites based upon several
factors: 1) previous mark–recapture history at each site (SAS,
unpubl. data); 2) permission from landowners for site accessibil-
ity (almost all Bog Turtle sites in Maryland are on private land);
and 3) estimated numbers of females at each site. Pressures posed
by the possibility of illegal collection led us to keep exact site
locations confidential; however, all selected sites are in the
piedmont of Maryland. The first site, ‘‘HA411’’, is a small (0.2 ha)
wetland situated between two roads and a stream. This site was
used previously by Morrow et al. (2001) for a study on Bog Turtle
movements. Parts of the site were actively grazed by sheep
during 2001, but, as of 2013, the site no longer is grazed. The site
is fed by a permanent spring and many seeps and is dominated
by sedge (Carex and Scirpus spp.), rushes (Juncus spp.), sensitive
fern (Onoclea sensibilis), and rice cut grass (Leersia oryzoides). There
is an aggregation of woody vegetation in the center of the
wetland, primarily in the form of smooth alder (Alnus serrulata).
The proximity of this wetland to a stream (5–10 m) causes the
southern portion floods regularly. This site has an even adult sex
ratio and a large Bog Turtle population, with 132 individual
turtles of both sexes captured and marked between 1992 and
2012 (SAS, unpubl. data).
The second site, ‘‘BA030’’, is a larger wetland (>1 ha) situated
in a developed suburban landscape. The most heavily surveyed
segment of habitat is situated between a power line right-of

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 HOME RANGES OF BOG TURTLES
way, a suburban backyard, a stream, and a drier hillside
dominated by deciduous vegetation. This primary survey area
has small patches of reed canary grass (Phalaris arudinacea) and
cattail (Typha latifolia) but otherwise is composed of rice cut
grass, sedges, and sensitive fern. Some of these patches of reed
canary grass have expanded in recent years, although these
patches are primarily along the edges of the wetland. The center
of the wetland has a large stand of smooth alder, and black
willow (Salix nigra) provides canopy cover around the
periphery of the habitat. Although most of the surveying and
mark–recapture effort at this site has been based on this focal
area, the overall wetland complex is patchily distributed to both
the south and the east of the focal survey habitat. Several
females have been caught in portions of the wetland immedi-
ately to the south of this focal site, and surveys were conducted
on that property as well. This site has fewer females than
HA411, with only 41 individual Bog Turtles of both sexes
captured in the past 20 years. The adult sex ratio of this site is
skewed in favor of females, with approximately four females for
every male (SAS, unpubl. data).
Initial Surveys.—We conducted Phase II and III Bog Turtle
surveys (as specified in Klemens, 2001) in April and May 2013
and 2014 to capture female Bog Turtles. Phase II surveys involved
one to three researchers opportunistically probing in shallow
pools, small mammal tunnels, tussock mats, and other areas
where Bog Turtles may hide in wetlands. Visual encounter
survey methods also were used when appropriate. Phase III Bog
Turtle surveys involved the use of passive entry traps and were
used only in 2014 to capture turtles at BA030. No additional
females were captured using this technique. We collected body
measurements (carapace and plastron length, body mass) from
each captured female and marked unmarked turtles using a
system modified from Ernst et al. (1974).
Radiotelemetry.—We captured female turtles at each site during
April and May in each study year, and attached SOPR-2190 radio
transmitters (Wildlife Materials, Murphysboro, IL) to each female
using methods outlined by Schubauer (1981) and Eckler et al.
(1990). Transmitters measured 33 · 13 · 8 mm and weighed 5–6 g
each. This is equivalent to 5–7% of adult body mass, which is well
below the 10% of body mass threshold acceptable for radiotelem-
etry of turtles (Guidelines for Use of Live Amphibians and Reptiles
in Field Research; http://www.aaalac.org/accreditation/
Guidelines_for_Use_of_Live_Amphibians_and_Reptiles.pdf). We
attached transmitters to female Bog Turtles with PC-7 epoxy.
After transmitter attachment, we monitored turtles overnight to
allow the epoxy time to dry. We then released captured turtles
within 24 h at the initial capture site (Eckler et al., 1990). We
collected radio locations twice a week in April and May, every day
during the peak nesting season of early June to early July (Ernst
and Lovich, 2009), and twice a month from late July to August.
Statistical Analyses.—We calculated 95% minimum convex
polygons (MCPs) using the ‘‘adehabitatHR’’ package in R version
3.0.3 (Calenge, 2006) and calculated fixed kernel estimates using
the program Geospatial Modeling Environment (GME; http://
www.spatialecology.com/). For kernel estimates, we first gener-
ated a probability distribution in GME using the Least Squares
Cross Validation (LSCV) bandwidth estimator, an algorithm that
can estimate kernel home-range sizes with the least bias when
compared to other kernel density bandwidth algorithms (Seaman
and Powell, 1996). Then, we created 50% and 95% isopleths.
Before analyzing home-range data, we tested for normality using
a Shapiro–Wilks test (Shapiro and Wilks, 1965) and tested for
homoscedasticity using Bartlett’s test (Bartlett, 1937). Home-
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69
range data violated both of these assumptions; however, a log
transformation corrected these violations.
Before analyzing home-range data, we excluded a random
study year for each female tracked in both years; in cases where
a female tracked during both years was tracked fewer than 10
times in a given study year, that study year was excluded for
that female. In this way, a data set composed only of unique
females captured in each site-by-year combination was created.
We used a two-way ANOVA with site and year as fixed effects
and 95% MCP home-range size as a response variable to test the
following null hypotheses: 1) year does not affect the 95% MCP
home-range size; and 2) site does not affect the 95% MCP home-
range size. Similar two-way ANOVAs were used to test the
effects of site and year on 50% and 95% KE home-range size. We
used a mixed-effect model to assess differences between home-
range estimators with type of estimator (95% MCP, 95% Kernel,
or 50% Kernel) as a within-subjects effect nested within turtle
identity and log-transformed home-range size as the response
variable. This model was created using the package ‘‘nlme’’ in R
(Pinheiro et al., 2013). We used a likelihood ratio test to compare
this model to a null model and a Tukey’s HSD multiple
comparison procedure using a Bonferroni correction to compare
pairwise differences between estimators using the package
‘‘multcomp’’ in R (Hothron et al., 2008).
We used a separate data set, composed only of females
tracked in both years and tracked >10 times in each study year
to investigate yearly variation in home-range size for females
tracked in both years. We used a mixed-effect model to assess
differences between years with year as a within-subjects factor
nested within turtle identity and log-transformed home-range
size as the response variable. A separate model was created for
each home-range estimator. We also created these models with
the package ‘‘nlme’’ in R (Pinheiro et al., 2013), and we used
likelihood ratio tests to compare these models to null models.
Unless otherwise noted, summary statistics are reported as
mean 6 SD: for all statistical analyses involving P-values, a =
0.05.
RESULTS
Differences in Home-Range Size between Study Years and
Locations.—We tracked 21 females during 2013, 11 at HA411,
and 10 at BA030. We tracked 25 females during 2014, 14 at
HA411, and 11 at BA030. We tracked 13 individual females in
both 2013 and 2014, with 7 at HA411 and 6 at BA030. For the data
set composed of unique females in each study year, final sample
sizes for BA030 were 8 for 2013 and 6 for 2014. For HA411,
sample sizes were 6 for 2013 and 12 for 2014. Turtles were tracked
an average of 19 times each season (range = 10–28). The highest
net daily movement distances (m/day) were in May (x = 0.378 6
0.535) and June (x = 1.32 6 1.83); the lowest were in April (x =
0.166 6 0.180) and August (x = 0.100 6 0.219).
Home-range sizes at BA030 typically were about twice larger
than home-range sizes at HA411 (Table 1). We found a
significant difference in home-range sizes between sites on
95% MCP estimates (F1,28 = 6.883, P = 0.0139) but did not find a
significant effect of year (F1,28 = 0.114, P = 0.738) or a significant
interaction (F1,28 = 0.484, P = 0.492). A similar test for 95% KE
home-range size found a significant difference in home-range
sizes between sites (F1,28 = 4.276, P = 0.048) but also did not
find a significant effect of year (F1,28 = 0.296, P = 0.591) or a
significant interaction (F1,28 = 0.482, P = 0.493). The ANOVA for
50% KE home-range size did not find a significant difference in
 70 N. W. BYER ET AL.
TABLE 1. Average (6 SD) home-range sizes for each site, year, and estimator, after random removal of one study year for females tracked both
years.
Site Year 95% MCP (m2)
BA030 2013 1,565.96 6 1,003.63
2014 1,708.10 6 1,513.36
HA411 2013 753.02 6 405.06
2014 820.45 6 518.92
home-range sizes between sites (F1,28 = 3.443, P = 0.074) or
between years (F1,28 = 0.204, P = 0.654), and did not find a
significant interaction (F1,28 = 0.950, P = 0.338). The analysis of
differences between years for females tracked both years did not
reveal significance differences between year for 95% MCP
(likelihood ratio = 0.19, P = 0.66), 95% KE (likelihood ratio =
0.007, P = 0.93) or 50% KE estimates (likelihood ratio = 0.0006,
P = 0.98).
Differences in Home-range Size between Estimators.—Typically,
95% KE estimates were 3–4 times larger than the 95% MCP
estimates, and 50% KE estimates were similar to 95% MCP
estimates (Table 1; Fig. 1). We found significant differences
between estimators (likelihood ratio = 48.87, P < 0.001). We
found a significant difference between 95% KE and 50% KE
estimates (z = 8.358, P < 0.001) and between 95% MCP and 95%
KE estimates (z = -6.760, P < 0.001) but did not find a significant
difference between 95% MCP and 50% KE estimates (z = 1.598, P
= 0.25).
DISCUSSION
Differences in Home-Range Size between Study Years and
Locations.—Chase et al. (1989) reported 95% MCP home-range
sizes of 2,400 m2 or less for Bog Turtles at two sites in Maryland,
whereas Morrow et al. (2001) reported 95% MCP home-range
sizes as high as 31,200 m2 at one of their two study sites (HA406).
FIG. 1. Box and whiskers plot of home-range sizes for the three
estimation techniques.
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95% KE (m2) 50% KE (m2)
5,680.88 6 6,896.28 1,248.41 6 1,725.78
6,265.20 6 5,580.79 1,384.62 6 1,094.02
2,961.39 6 1,438.33 688.58 6 315.94
2,860 6 1,923.71 660.01 6 488.05
Morrow et al. (2001) also used HA411 as one of their study sites;
they reported average 95% MCP home-range sizes of 200 m2 for
1996 and 300 m2 for 1997. These are much smaller than the
average 95% MCP home-range sizes documented for HA411
during our study (753 m2 for 2013 and 820 m2 for 2014). Morrow
et al. (2001) also documented an increase in average home-range
sizes over time at another Bog Turtle site in Maryland, and
suggested this may reflect a decrease in site quality.
For HA411, the increase in home-range sizes from 1996–1997
to the present day also may reflect a decrease in habitat quality.
Woody invasive species including wineberry (Rubus phoenicola-
sius) have proliferated at this wetland in recent years. Wineberry
was not documented on species lists presented in Morrow et al.
(2001) and may represent a recently established invasive species
at this site. Native woody shrubs such as smooth alder, swamp
rose, and black willow appear to dominate the central portion of
HA411 (pers. obs.). These woody plants may out-compete
herbaceous native vegetation preferred for nesting (such as
tussock sedge) or hiding (such as sensitive fern). In addition,
although data for our study were collected between the months
of April and August as in Morrow et al. (2001), we collected a
large majority of the telemetry data in June, during the nesting
season. Calculated daily movement distances were higher in
June than in any other month, suggesting that either the
frequency of data collection or nesting-related movements may
have affected telemetry data in our study. This difference in
radiotelemetry effort may account for observed differences in
home-range sizes between our study and Morrow et al. (2001),
although proliferation of woody and invasive vegetation is
another potential cause.
We documented significant differences in 95% MCP and 95%
KE home-range sizes between BA030 and HA411 but no
significant differences in 50% KE home-range sizes between
BA030 and HA411. This could suggest that core used areas
(approximated by 50% KEs) may be relatively similar for Bog
Turtles at each site. Average 50% KEs at BA030 were almost
twice as large as those at HA411, however, despite the lack of
statistical significance for this comparison; in addition, 50% KEs
at each site had very high standard deviations, possibly
accounting for the nonsignificant difference between the two
sites. With these caveats, all estimated home-range sizes
appeared larger for BA030 than for HA411. BA030 is a larger
wetland than HA411. Because Bog Turtles spend most of their
time within wetland boundaries (Ernst and Lovich, 2009), this
may partially explain the higher home-range sizes at BA030 and
would suggest that turtles in larger wetlands should typically
move longer distances to take advantage of available high-
quality habitat. However, reed canary grass has proliferated at
BA030 since initial surveys were conducted in 1992 (SAS, pers.
obs.). Reed canary grass is invasive throughout North America
and can form dense monocultures within wetlands (Lavergne
and Molofsky, 2004). In Bog Turtle wetlands, reed canary grass
and other invasive plants reduce light penetration to the
wetland surface and out-compete native plants that Bog Turtles
 HOME RANGES OF BOG TURTLES
may need for foraging, nesting, and basking (Morrow et al.,
2001). If wetland quality is deteriorating at this site attributable
to the spread of reed canary grass, Bog Turtles would need more
habitat area to fulfill habitat requirements (Morrow et al., 2001).
Differences in Home-range Size between Estimators.—Although no
statistically significant difference was detected between 50% KEs
and 95% MCPs, 95% KEs were significantly larger than 50% KEs
and 95% MCPs by a factor of six or more. For this study, we used
a relatively well-established smoothing parameter algorithm
(LSCV). We chose this smoothing parameter because very few
published studies have used kernel estimates for Bog Turtles,
with the exception of Morrow et al. (2001); but Morrow et al.
(2001) did not specify their choice of smoothing parameters.
Although Row and Blouin-Demers (2006) suggested that kernel
estimates using the LSCV bandwidth estimator are unreliable for
herpetofauna, we wanted to evaluate this common bandwidth
estimator for this species before evaluating alternatives to this
estimator, because Bog Turtles typically have much smaller
home-range sizes than other reptiles.
Our study supports the assertions of Row and Blouin-Demers
(2006) about the unreliability of kernel estimates for herpeto-
faunal field studies, because 95% kernel estimates varied
considerably, and typically were much larger than 95%
minimum convex polygons. Surprisingly, 95% minimum convex
polygons and 50% kernel estimates with the LSCV bandwidth
estimator produced similar home-range sizes in our study. We
urge caution in interpreting these patterns in home-range sizes,
however, because the number of telemetry locations per female
in this study was small. Row and Blouin-Demers (2006)
suggested that the number of telemetry locations will alter
kernel home-range sizes, indicating that relatively small sample
size may have inflated kernel home-range sizes in this study.
Lanver and Kelly (2008) suggested that unified and estab-
lished standards should be developed for each field regarding
the reporting and methodology of home-range estimates. Our
study suggests the methods outlined in Row and Blouin-
Demers (2006) represent a solid foundation for evaluating
home-range estimator choices for reptiles or at least for turtles.
Because this group encompasses many different body plans,
natural histories, and life-history strategies, however, research-
ers should recognize that developing general practices for all
reptiles may be difficult. Other studies have documented
species and taxa-specific patterns of differences between
home-range estimators (Barg et al., 2005; Huck et al., 2008).
Therefore, the choice of a home-range estimator should be based
upon study design and study species, because both of these
factors can influence home-range estimates (Börger et al., 2006;
Boyle et al., 2009).
Conservation Implications.—To incorporate home-range sizes
into habitat management efforts for this species, biologists must
collect telemetry data on a site-specific basis and should take site
area into account when designing studies. In addition, home-
range estimators must be chosen carefully, as considerable
variation in home-range sizes is apparent when using different
estimation techniques. Habitat management efforts should also
collect information on changes in home-range size over time, as
such information may be useful for assessing responses of turtles
to habitat modification. Although our study did not document an
effect of year on home-range size, the observed increase in home-
range size at HA411 since Morrow et al. (2001) may indicate that
changes in home-range size over time are mainly driven by
relatively slow changes in wetland vegetation, specifically woody
species. These changes may not be detected by studies that collect
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71
home-range data over two or three consecutive years. Therefore,
management efforts that incorporate radiotelemetry may need to
collect home-range information at study sites every 5–10 years if
researchers hope to detect changes in home-range size and
habitat quality. By incorporating these considerations, conserva-
tion efforts could more accurately estimate patterns of habitat use
for this elusive species.
Acknowledgments.—Funding for this study was provided by
the U.S. Fish and Wildlife Service, the Maryland Department of
Natural Resources, and the Towson University Department of
Biological Sciences. This study was approved by the Towson
University IACUC committee (IACUC FR 021913 RS-04) and
the Maryland Department of Natural Resources (permit 53959).
We thank A. Byer, K. Anderson, R. McGehee, S. Martin, S.
McDaniel, and many others for their help and guidance with
fieldwork and study design. We also thank our site owners,
without whom this work would not have been possible. S.
Martin and A. Byer provided comments and suggestions on the
manuscript.
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Accepted: 24 June 2016.
Published online: 11 January 2017