2012 NORTHEASTERN NATURALIST 19(3):361–372
Notes on the Nesting Ecology of Eastern Box Turtles near
the Northern Limit of their Range
Lisabeth L. Willey1,2,* and Paul R. Sievert2
Abstract - We evaluated the nesting ecology of Terrapene carolina carolina (Eastern
Box Turtle) near their northern range limit in Massachusetts. We identifi ed 34 nests in
2005 and 2006 at 4 study sites, and measured clutch size, nest success, hatchling size, and
habitat characteristics at each site. Mean clutch size was 5.87 eggs, and egg survival was
approximately 50% to hatching, excluding depredation. Large-bodied females tended to
oviposit larger clutches than small-bodied females, although the correlation was not signifi
cant, and a smaller proportion of their eggs produced live hatchlings. Nest depredation
varied greatly across sites from 0 to nearly 100%. The variability observed across the
species’ range and across sites underscores the importance of obtaining local information
when developing conservation and management programs for rare turtles. The characteristics
of the nest sites observed in our study could be simulated to more effectively create
or maintain artifi cial nest sites for Eastern Box Turtles in the Northeast.
Although Terrapene carolina carolina L. (Eastern Box Turtle) is well studied
throughout much of its range, few studies have examined the species in New
England, where it occurs in relatively low densities. Box turtles are protected
in most northeastern states in which they occur. In Massachusetts, Eastern Box
Turtle is protected as a species of special concern under provisions of the Massachusetts
Endangered Species Act (MESA) (M.G.L. Ch. 131A), and the state
is currently undergoing a comprehensive conservation planning process for the
species (Erb 2011). To inform this and other regional conservation efforts, we
evaluated the nesting ecology of Eastern Box Turtles near the northern limit of
the species’ range.
Eastern Box Turtle nesting ecology has been examined at numerous sites
throughout the species’ range (e.g., Allard 1935, Burke and Capitano 2011, Flitz and
Mullin 2006, Kipp 2003, Wilson and Ernst 2005), and based on the hypothesis proposed
by Iverson (1992) that clutch size increases with latitude, we expected to see
larger clutch sizes than those observed in these more southerly studies. Recent work
on the relationships among geographic location within a species’ range, abundance,
and demographic parameters (e.g., Angert 2009, Gerst et al. 2011) demonstrate the
complexity of these relationships, and underscore the importance of assessing variability
across species’ ranges and obtaining local demographic parameter values to
develop more informed and effective conservation strategies.
1Graduate Program in Organismic and Evolutionary Biology, University of Massachusetts
Amherst, North Pleasant Street, Amherst, MA 01003. 2United States Geological
Service, Massachusetts Cooperative fish and Wildlife Research Unit, Department of Environmental
Conservation, University of Massachusetts Amherst, 160 Holdsworth Way,
Amherst, MA 01003. *Corresponding author - firstname.lastname@example.org.
362 Northeastern Naturalist Vol. 19, No. 3
Availability of nesting habitat with a suitable thermal regime is one potential
factor limiting the distribution of Eastern Box Turtles and other non-marine turtle
species (Allard 1935, Bobyn and Brooks 1994, Compton 1999). To address this
potential limitation, anthropogenic improvement or creation of nesting habitat
for rare turtles is commonly conducted throughout the Northeast (Beaudry et al.
2010, Kiviat et al. 2000, MNHESP 2009), but the nesting habitat requirements
for Eastern Box Turtles in the region have not previously been explored. This
information is important for developing such site-specifi c management plans and
to assist the state-wide conservation planning process.
We assessed nesting habitat, clutch size, and nest success by radio-tracking
female box turtles at 4 study sites in 2005 and 2006. We also related clutch size
and nest success to female body size and habitat characteristics.
This study was conducted at 4 study sites in the Connecticut River Valley,
MA. Study sites were selected across a range of human land-use intensity, habitat
type, latitude, and elevation (Table 1). Sites were distributed throughout the Valley
along a north–south gradient of 56 km and ranged in elevation from 50 m to
300 m. The use of multiple sites with different habitat characteristics provides an
opportunity to estimate variation in nesting parameters. To quantify habitat differences
between sites, the percentage of forested, open canopy, and developed
area within each site was measured using 1999 land-use cover data from the Offi
ce of Geographic and Environmental Information (MassGIS), Commonwealth
of Massachusetts Executive Offi ce of Energy and Environmental Affairs. The 4
sites are described below. The exact names and localities of the sites are withheld
for conservation purposes.
Site A (50 ha) is a small, municipally owned conservation area that consists of
Acer rubrum L. (Red Maple)-dominated forest, a wetland complex, and a former
gravel-extraction site. This site is surrounded by active agriculture and residential
development and is heavily used by hikers.
Site B (100 ha) is a parcel of private conservation land consisting of a pineoak
upland and a beaver-dammed brook flowing through the low-lying portion
of the site. The adjacent lot was developed in 2004–2005, and onsite mitigation
Table 1. Summary of Eastern Box Turtle study sites.
Site Area % % open %
name (ha)A forestedB Dominant tree species habitatB developedB
Site A 50 25% Acer rubrum 32% 43%
Site B 100 78% Pinus strobus, Quercus spp. 11% 11%
Site C 900 79% Pinus rigida, Quercus spp., 16% 5%
Acer saccharum Marshall (Sugar Maple)
Site D 1200 90% Quercus species, Carya spp., Tsuga 5% 4%
canadensis (L.) Carrière (Eastern Hemlock)
ACalculated as the continuous land area to nearest paved road, used as an approximation for human
BCalculated as percent of area in specifi ed land use using MassGIS 1999 land-use data.
2012 L.L. Willey and P.R. Sievert 363
for box turtles was required under MESA. The development was surrounded by
a “turtle curb”, which consisted of a continuous concrete barrier greater than
30 cm in height, designed to confi ne the animals to the undeveloped area. An
artifi cial nesting area was also constructed within the Eastern Box Turtle habitat.
Within the last 20 years, the site has been surrounded by development including
residential neighborhoods, schools, and light industrial uses, primarily shipping
Site C (900 ha) is a state-owned conservation area where prescribed burns are
used for restoration and habitat maintenance. The site is used by the public for
recreation (e.g., hunting, walking, jogging, and all-terrain vehicles), and consists
of a deciduous forested hillside adjacent to a large tract of Pinus rigida Mill.
(Pitch Pine)-scrub oak barrens with areas of open sand.
Site D (1200 ha) comprises the south slope of an east–west-trending basaltic
mountain range (up to about 300 m in elevation). This site has little human use
(hiking, hunting, and some all-terrain vehicle use), and it is bordered by residential
development, a major state highway, and gravel operations to the east, south,
and west. This site is primarily composed of private forest land, but is bisected
by an east–west treeless right-of-way.
Turtles were captured using visual encounter surveys. We individually marked
each turtle by fi ling the marginal scutes (Ernst et al. 1974). We then measured each
animal using dial calipers, and photographed it in the fi eld. For each turtle, we measured
straight carapace length (SCL), total carapace length (TCL), plastron width
at the humeral/pectoral seam (PW), carapace width at the widest point (CW), and
carapace height at the deepest point (CH). We categorized turtles into 6 age classes
by counting the lines of arrested growth (LAG) on their plastron and evaluating
relative amounts of shell-wear on the plastron: Class 1 = visible new growth on
plastron, no wear; Class 2 = no new visible growth, no wear; Class 3 = beginning
to wear; Class 4 = less than 50% worn; Class 5 = more than 50% worn; Class 6 =
plastron worn smooth. At each site, a subset of turtles (5–11 adult females) was
outfi tted with radio transmitters (MBFT-6, Lotek Wireless, Newmarket, ON,
Canada; R2020, Advanced Telemetry Systems, Isanti, MN). We affi xed radios
along the posterior margin of the carapace using dental acrylic (Biocryl Resin,
Great Lakes Orthodotics, Tonawanda, NY). For MBFT-6 models, antennae were
affi xed with dental acrylic to the carapace in a ring along the pleural/marginal seam
(as in Compton 1999); for R2020 models, the antennae were shorter and were free
to trail behind the turtle. Total weight of radio and acrylic was less than 5% of body
weight. Turtles were released immediately after processing.
We located turtles using radio-telemetry 2–3 times per week throughout the
spring and May-to-June nesting season and recorded positions to the nearest 5 m
using a hand-held global positioning system (GPS) receiver (eTrex 12-channel or
GPSmap 76CSx, Garmin International, Inc., Olathe, KS). As radio-equipped females
neared potential nest sites, we located them more frequently (up to twice per
364 Northeastern Naturalist Vol. 19, No. 3
day) and weighed and palpated them to determine if they were gravid. If females
were gravid, we affi xed a thread bobbin to the caudal portion of the carapace (using
the method described in Milam 1997). When we located the animal in the morning,
we assessed whether she remained gravid. If she was no longer gravid, or if we
were unable to palpate her but she had lost considerable weight from the previous
night, we followed her thread trail to locate the nest chamber (Milam 1997). Potential
nest sites were indicated by an erratic thread trail, disturbed soil, or areas where
the thread was buried. We collected the thread and removed it from the site when
we located the chamber, or if it could not be located after substantial searching.
We covered nests with half-inch-mesh hardware-cloth screens to protect eggs
from predators. We replaced screens with hardware-cloth box screens in August
to allow hatchlings to emerge without escaping (Graham 1997), and later checked
nests daily for emerged hatchlings. We measured and released the hatchlings upon
emergence. Nests were excavated in November, after hatchlings had emerged, in
order to determine clutch size and nest success. Success was measured as the proportion
of eggs that developed and successfully emerged from the nest.
At each nest site, we collected environmental data, including cover type,
substrate type, distance to nearest ecotone, and vegetation structure within a 5-m
radius of the nest. Within that circle, we visually approximated percent canopy
(greater than 3 m in height), percent shrub cover (woody stems less than 3 m in
height), percent herbaceous cover, leaf litter, and bare ground. We also recorded
the 3 dominant plant species of each layer.
We measured the dimensions of the canopy openings used by turtles for nesting
using 2005 orthophotos (MassGIS) and ArcMap 9.2 (ESRI, Redlands CA).
We also measured the area of the canopy opening and the distance of each nest
to the nearest forest edge in the four cardinal directions via GIS.
We evaluated the effect of site and calendar year on clutch size and nest success
using ANOVA. We used linear regression to explore the effects of age, body
size, and habitat characteristics on clutch size and nest success. To maintain
independence of samples, only one nest per female was included in the analysis.
The fi rst (2005) nest of females with nests in both years was used. This procedure
resulted in a sample size of 24 for ANOVA and regression analyses. Count
values (i.e., age class, clutch size, and number hatched) were square root transformed,
and proportions (i.e., forest cover and nest success rate) were arcsine
transformed. Residuals were visually assessed to ensure assumptions were not
violated. Analyses were conducted in R (R development core team 2010).
We screened 34 Eastern Box Turtle nests at 4 sites in 2005 and 2006. Nesting
was observed in the Connecticut River Valley from 27 May to 10 July, with
the peak occurring in early June. The fewest LAGs exhibited on any female
that we observed nesting was 14 LAGs, and her SCL was 144 mm. We tracked
3 females with 12 and 13 LAGs (119–140 mm SCL), but these animals did not
nest. The smallest-bodied female that we observed nesting was 122 mm SCL,
but her carapace was entirely worn smooth, and she was probably very old.
2012 L.L. Willey and P.R. Sievert 365
Approximately 90% of mature females nested annually. Turtles were carefully
monitored for evidence of double clutching, and no instances were documented.
Of the 34 nests, two were depredated despite protective screens, and one was
destroyed by mechanized equipment. Results from the remaining 31 nests are
presented in Table 2. Clutch size ranged from 3 to 10 (mean = 5.87, sd = 1.88).
Twenty-four of the 31 nests produced at least one hatchling, and mean nest success
(average proportion of eggs hatching in each nest), excluding depredation
or human disturbance, was 53%. Of the 182 eggs oviposited in all 31 nests,
55% hatched successfully, 28% failed to develop, 16% died prior to hatching,
and 1% hatched but died in the nest chamber.
In 2005 and 2006, hatchlings emerged between 20 August and 9 October.
Upon emergence, hatchling carapace length ranged from 26.7 to 35.2 mm (mean
= 31.9, n = 48). Average hatchling size was not correlated with the body size of
the mother (F1,12 = 0.00, P = 0.94). The hatchlings from larger clutches tended to
be smaller in size, but not signifi cantly (F1,12 = 2.4, P = 0.14).
Correlation between body size, age, habitat and reproductive rates
Between sites, there were no signifi cant differences in clutch size (F3,20 = 1.3,
P = 0.31), hatching number (F3,20 = 0.94, P = 0.44), or success rate (F3,20 = 1.4, P =
0.26). Similarly, there were no signifi cant differences between years (F3,20 < 0.14,
P > 0.7 in all cases), and therefore sites and years were pooled for the remaining
analyses. Female body size was positively correlated with the square root of
clutch size (fig. 1A), and TCL was the body size metric best explaining clutch size,
although this relationship was not signifi cant (F1,22 = 1.98, P = 0.17). Carapace
height showed no linear relationship with transformed clutch size (F1,22 = 0.00,
P = 0.95); the three females that deposited the largest clutches (9 or 10 eggs) had
carapace heights (68–71 mm) close to the mean (68 mm) (fig. 1B). Transformed
nest hatching rate was negatively correlated with total carapace length (F1,22 =
5.0, P = 0.04) and carapace height (F1,22 = 4.5, P = 0.04). Larger females had more
frequent nest failures than smaller-bodied females (fig. 1D, E), but because larger
females oviposit larger clutches, the total number of successful hatchlings was
not signifi cantly correlated with the body size of the mother (F1,22 = 2.6, P = 0.12).
The mother’s age class, ranging from 1 to 6, had no signifi cant linear relationship
with transformed clutch size (F1,22 = 1.07, P = 0.31) or nest success (F1,22 = 0.88, P =
0.36) (fig. 1C, F).
Clutch size and nest success were not signifi cantly correlated with any habitat
variable (F1,22 < 1.47, P > 0.23 in all cases). Turtles in heavily forested areas
Table 2. Summary statistics for the size and success rate of 31 box turtle nests observed in the Connecticut
River Valley, MA. Analysis excludes nests that were depredated or destroyed.
Clutch size Successfully emerged hatchlings Success rate
Minimum 3 0 0.00
Median 6 3 0.67
Mean 5.87 3.23 0.53
Maximum 10 8 1.00
Standard deviation 1.88 2.49 0.35
366 Northeastern Naturalist Vol. 19, No. 3
tended to lay smaller clutches, whereas those in more open habitats deposited
larger, less successful clutches, although these relationships were not signifi cant
(F1,22 = 1.47, P = 0.24).
Although we screened nests of radio-tagged turtles and therefore did not
directly measure the effects of depredation on box turtle nests, 2–6 incidentally
observed box turtle nests at each site were not screened. The effect of depredation
on nests that were not screened appeared to vary considerably. Although sample
size was low, sites with higher nest density and a greater number of turtle species
present that were located in areas surrounded by anthropogenic land cover had
higher depredation rates (up to 100%). All 4 nests were completely destroyed by
predators at the site (Site A) that is surrounded by residential and agricultural
land uses. Two of these nests were screened and 2 were not. Six unscreened nests
located in a remote power line right-of-way through a forested area (Site D) were
left completely intact (0% depredation).
Turtles used abandoned gravel pits, right-of-ways, backyards, old fields,
and forest clearings as nest sites. Nest sites tended to be sandy, open areas
with little vegetation, but all nests were deposited within 1 m of vegetation.
figure 1. Eastern Box Turtle clutch size and nest success plotted against body size. Larger
females tend to lay larger clutches (though not signifi cantly), and they are signifi cantly
less successful than those oviposited by smaller bodied females.
2012 L.L. Willey and P.R. Sievert 367
Canopy cover within 5 m of nests (n = 34) ranged from 0 to 65% (median = 0),
shrub cover within 5 m ranged from 0 to 40% (median = 5%), and herbaceous
cover within 5 m ranged from 5% to 90% (median = 45%). Quercus spp. (oak)
dominated the canopy and shrub layers whereas various species of graminoids,
Solidago spp. (goldenrod), and Potentilla spp. (cinquefoil) were the most commonly
occurring herbaceous plants around the nest.
At nest sites, turtles spent more time near the nest area if woody or herbaceous
material in which to hide and forage was present. Areas with downed wood,
graminoids, goldenrod species, Comptonia peregrina (L.) Coult. (Sweet Fern),
Rubus spp. (blackberry/raspberry/dewberry), Rhus typhina L. (Staghorn Sumac),
Rosa multiflora Thunberg (Multiflora Rose), Lonicera spp. (Honeysuckle), and
Elaeagnus umbellata Thunberg (Autumn Olive), as well as young Betula spp.
(birch), Populus spp. (aspen), and oak species were frequently used for resting
and foraging by females immediately before and after nesting.
Nest-site open-canopy dimensions
There were 16 nesting areas at the 4 study sites. Median dimensions of clearings
used by nesting turtles were 125 m on the north–south axis and 54 m on the
east–west axis. Nest site openings were generally longer on the north–south axis.
Minimum axes were 19 and 7 m, respectively. The minimum observed opening
used as a nest site was approximately 1200 m2, and the median was 5670 m2
(Table 3). Table 4 lists the distances of observed turtle nests from various forest
edges. All of the nests we observed were closer to the northern edge of sandy
openings than to the southern edge.
Table 3. Dimensions of canopy openings used by Eastern Box Turtles for nesting.
Dimension Minimum Median Maximum
North–south axis length (m) 19 125 1000A
East–west axis length (m) 7 54 377
Approximate clearing area (m2) 1200 5670 72,384
AFor clearings that are very long (e.g., power-line corridors), a maximum dimension of 1000 m
Table 4. Distances (m) from Eastern Box Turtle nests to forest edges.
Minimum 5th percentile Median Maximum
North 0 0.0 12 500A
Northeast 0 3.2 16 100
East 0 2.4 27 78
Southeast 14 14.0 35 117
South 10 22.0 125 500A
Southwest 12 15.2 43 175
West 0 3.2 27 361
Northwest 0 0.8 20 52
AFor clearings that are very long (e.g., power-line corridors), a maximum distance of 500 m was used.
368 Northeastern Naturalist Vol. 19, No. 3
Clutch size and nest success
The clutch size we observed was larger than that reported from most other
studies (Table 5). There was no evidence of double clutching in our sample, although
this has been reported fairly regularly farther south; double clutching was
also not observed in Long Island (Cook 2004). Our results support the observation,
generally suggested in the literature (e.g., Iverson 1992, Iverson et al. 1993,
Wilson and Ernst 2005) that clutch size increases with latitude, whereas clutch
frequency decreases with latitude. The proportion of females nesting in a year
(90% in our sample) was higher than that observed in other studies (Cook 2004,
Dodd 2001, Wilson and Ernst 2005).
Nest success rate (55%), which excluded depredation, was similar to that
reported in other studies (e.g., Belzer 2002, Kipp 2003, Wilson and Ernst 2005).
However, the methodology among studies in estimating success varied, so it is
diffi cult to directly compare predation rates. While it is possible that human intervention
(i.e., fi nding and screening the nest) lowered success rates, our methods
were similar and in many cases less invasive than those used in other studies. In
addition, Samson et al. (2007) found that handling did not decrease the rate of
nest productivity in Chrysemys spp. (painted turtles), suggesting a natural mechanism
for the low success observed in our case.
Because nesting habitat with the appropriate thermal regime for successful
incubation is thought to be one factor limiting turtles at the northern limit of
their range (Allard 1935, Bobyn and Brooks 1994, Compton 1999), low success
rates are not surprising and suggest that any advantage conferred by larger clutch
sizes in northern areas may offset lower success rates. Limited nesting habitat
availability and incubation temperature suitability in Massachusetts may explain
why box turtles tend to be distributed in the warmer, sandier portions of the state:
Cape Cod, the southeast, and the Connecticut River Valley. Future work on the
thermal regime of nests near the northern edge of the species range may help
elucidate this question.
Variation in both clutch size and success rate across the species’ range underscores
the importance of adjusting parameters to local demographics when
Table 5. Clutch sizes of Eastern Box Turtles reported throughout their range.
Author Location Average clutch size
Allard 1935 Washington, DC 4.2
Ewing 1935 Washington, DC 3.8
Congdon and Gibbons 1985 South Carolina 3.4
Stuart and Miller 1987 North Carolina 3.0
Mitchell 1994 Virginia 4.1
Tucker 1999 Illinois 4.9
Belzer 2002 Pennsylvania 4.0
Kipp 2003 Delaware 4.6
Cook 2004 Long Island, NY 5.8
Wilson and Ernst 2005 Virginia 3.15
Burke and Capitano 2011 Long Island, NY 4.1
This study Massachusetts 5.87
2012 L.L. Willey and P.R. Sievert 369
assessing status and developing conservation plans. For instance, a populationviability
analysis utilizing the larger northern clutch size with a higher observed
nest success rate from another regional study would be positively biased with regard
to recruitment and population growth. Conversely, using smaller clutch sizes
reported for more southern populations with mortality rates from a site with lower
nest success rates would lead to exaggerated projections of population decline.
Although the positive correlation between clutch size and body size has been
reported in a number of studies (Iverson 1992, Kipp 2003, Tucker 1999, Wilson
and Ernst 2005), few studies have evaluated the correlation between the mother’s
body size and number of successful hatchlings. Our results suggest that in clutches
deposited by larger mothers, a smaller proportion of eggs successfully hatch,
potentially offsetting the benefi t conferred by larger clutch size, although larger
clutches still produced signifi cantly more hatchlings (F1,22 = 4.62, P = 0.04).
Although smaller females tended to produce a greater proportion of successful
eggs, because there was no correlation between female body size and the total
number of hatchlings or hatchling size, and because we did not follow hatchlings
beyond emergence, we do not know the associated recruitment rates or whether
larger- or smaller-bodied turtles are more likely to produce offspring that survive
Allard (1935) and Tucker (1999) found similar results whereby large female
Eastern Box Turtles deposited larger clutches, but large clutches contained eggs
that were smaller and weighed less, generally agreeing with the pattern discussed
by Iverson et al. (1993). We did not measure or weigh eggs in our sample, so we do
not know whether there was a correlation between egg size and hatching rate.
High depredation rates of turtle nests are frequently reported. For example,
in Illinois, Flitz and Mullin (2006) observed 87.5% depredation on Eastern
Box Turtle nests that were not protected. However, predation rates are difficult
to measure because they vary substantially across both space and time
with weather, predator densities, landscape characteristics, etc. (e.g., Bowen
and Janzen 2005, Kolbe and Janzen 2002, Strickland and Janzen 2010), and it
is possible that observer disturbance increases susceptibility to nest predation
(Rollinson and Brooks 2007). We found that depredation rates varied greatly by
location, similar to variation previously reported in Eastern Box Turtles (e.g.,
Kipp 2003). While our sample size was not large enough to draw statistical conclusions
about the relationship between habitat variables and nest depredation
rates, depredation rates on nests are likely a function of local predator density
as well as turtle nest density. In other systems, edges have been shown to increase
the prevalence of meso-predators (Dijak and Thompson 2000, Herkert
et al. 2003), and in some cases, turtle nests near edges have been shown to be
more susceptible to depredation (Strickland and Janzen 2010, Temple 1987),
but this effect has not been consistent across studies (Kolbe and Janzen 2002,
Marchand and Litvaitis 2004).
370 Northeastern Naturalist Vol. 19, No. 3
The 3 nests of the original 34 nests that were destroyed by predators and humans
were located at sites with the greatest amount of human activity. Both sites
were surrounded by residential development. Site A, the site with depredation rates
up to 100%, is a residential and agricultural area that may support larger numbers
of meso-carnivores. Chrysemys picta picta Schneider (Eastern Painted Turtle) and
Chelydra serpentina serpentina L. (Eastern Snapping Turtle) use the same nesting
area as Eastern Box Turtles at this site, creating turtle nest densities that were higher
than those at other sites (L.L. Willey, pers. observ.). Site D, which had very low
depredation, is mostly forested, and no other species of turtle has been observed
there. While limited, our observations seem to support the conclusion of previous
authors (e.g., Marchand and Litvaitis 2004) who observed that depredation was
higher in areas with high nest density and near agricultural sites.
Habitat management implications
Clearing land to open up habitats for nesting turtles is conducted throughout
Massachusetts by both MassWildlife and USDA Natural Resources Conservation
Service (MNHESP 2009; B. Schreier, USDA Natural Resources Conservation
Service, Amherst, MA, pers. comm.) and throughout the Northeast (Kiviat et
al. 2000). Our results can be used to inform the design of such management and
suggest that canopy openings should be at least 1200 m2 and probably larger to
attract nesting box turtles. Openings where turtles nested were generally longer
on the north–south axis than the east–west axis. This fi nding is probably related
to insolation. Interestingly, females that inhabited east–west oriented powerline
habitat prior to nesting in May sometimes traveled up to 1200 m to nest
elsewhere, whereas those that used north–south oriented power-lines generally
nested at that location. Sample size was small, so other attributes (e.g., soil or
vegetation type) might have influenced nest-site choice. We would still recommend
that nesting habitat should be created on a north–south axis.
Beaudry et al. (2010) reported that adult Emydoidea blandingii Holbrook
(Blanding’s Turtle) are willing to use anthropogenically altered areas, but there
is no empirical evidence to evaluate nesting success in such sites. Future studies
should address the frequency and viability of nests in managed areas. Ideally,
management plans should be developed with all life stages in mind, but particular
precaution should be taken to avoid adult mortality during management activities.
We thank the Massachusetts Natural Heritage and Endangered Species Program, The
University of Massachusetts Graduate Program in Organismic and Evolutionary Biology,
the Department of Natural Resources Conservation, the USGS Massachusetts Cooperative
fish and Wildlife Research Unit, The University of Massachusetts Natural History
Collections, and the Turtle Conservation Project for funding support. We also thank M.
Jones, L. Johnson, Z. Dowling, D. Yorks, C. Jordan, B. Dunphy, and B. Crowley for their
help in the fi eld, and A. Breisch, R. Cook, S. Fowle, C. Griffi n, M. Jones, K. McGargial,
A. Richmond, and two anonymous reviewers for providing helpful comments on previous
versions of this manuscript. We thank the many landowners for use of their property
during the course of this study. Methods were approved by the University of Massachusetts
at Amherst Institutional Animal Care and Use Committee (protocol # 25-02-04).
2012 L.L. Willey and P.R. Sievert 371
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