The Journal of Wildlife Management 78(1):101–111; 2014; DOI: 10.1002/jwmg.

645

Habitat Relations

Effects of Body Size and Sex of Drymarchon

couperi (Eastern Indigo Snake) on Habitat
Use, Movements, and Home Range Size
in Georgia
NATALIE L. HYSLOP,1,2 Warnell School of Forestry and Natural Resources, The University of Georgia, Athens, GA 30602-2152, USA
J. MICHAEL MEYERS, USGS Patuxent Wildlife Research Center, Warnell School of Forestry and Natural Resources, The University of Georgia,
Athens, GA 30602-2152, USA
ROBERT J. COOPER, Warnell School of Forestry and Natural Resources, The University of Georgia, Athens, GA 30602-2152, USA
DIRK J. STEVENSON,3 Fort Stewart Directorate of Public Works Fish and Wildlife Branch, United States Army, Fort Stewart, GA 31314, USA

ABSTRACT The federally threatened eastern indigo snake (Drymarchon couperi), native to the southeastern
Coastal Plain of the United States, has experienced population declines caused primarily by habitat loss,
degradation, and fragmentation. To examine spatial and habitat use requirements of the species, we
radiotracked 32 eastern indigo snakes from 2002 to 2004 on Fort Stewart Military Installation and adjacent
private lands in Georgia. We estimated annual and seasonal home ranges and evaluated a priori hypotheses
examining morphometric and ecological factors (sex, body size, location) associated with intraspecific
differences in home range size. We analyzed habitat use hierarchically by examining use across the study area
and within home ranges. Annual home range size varied from 33 ha to 1,528 ha (average minimum convex
polygon: x2003 ¼ 378 ha; x2004 ¼ 340 ha). Individual home range size was most influenced by sex (males with
larger home ranges) followed by body size. Compositional analysis of habitat use suggested positive selection
for wetland, evergreen forest, and pine-hardwood (mixed) forest, with an avoidance of roads and deciduous
forests. Seasonally, indigo snakes used the highest diversity of habitats as they moved from xeric uplands
(sandhills) in winter and early spring to wetlands and uplands other than sandhills in summer; however,
snakes continued to use sandhill habitats (35–58% of locations seasonally) with gopher tortoise (Gopherus
polyphemus) burrows throughout the warmest months. In Georgia, management and conservation of the
eastern indigo snake should include conservation of large tracts of undeveloped land, containing a matrix of
xeric uplands with suitable underground shelters and adjacent wetland habitats. Ó 2013 The Wildlife
Society.

KEY WORDS conservation, Drymarchon couperi, eastern indigo snake, Georgia, gopher tortoise, habitat use, home
range, movement, telemetry.

Accurate natural history information, including the spatial spatial arrangement, connectivity, and availability of necessary
and habitat requirements of a species, is necessary for resources such as prey, suitable shelters, thermoregulatory
development of adaptive conservation and management plans. opportunities, and mates (Gibbons and Semlitsch 1987,
These needs emphasize the importance of understanding Gregory et al. 1987, Macartney et al. 1988). Understanding
habitat components and spatial requirements for maintaining the scale at which populations are structured and determining
populations across their geographic range. Spatial patterns of factors associated with home range size and habitat use are
population structure can often be understood by examining important for management strategies and the design of
the distribution of individuals in relation to required resources adequately sized reserves (Dodd 1987).
(Pope et al. 2000). Possible influential factors include the In the southeastern Coastal Plain, a center of reptile and
amphibian diversity in the United States, habitat loss is the
primary cause of most herpetofaunal declines (Gibbons
Received: 8 March 2012; Accepted: 22 September 2013 et al. 1997). Contributing to these declines has been the loss of
Published: 5 December 2013 the once dominant longleaf pine (Pinus palustris)–wiregrass
1
(Aristida stricta) habitats, which historically covered approxi-
E-mail: nhyslop@ung.edu mately 30 million ha of the southeastern Coastal Plain prior to
2
Present address: Department of Biology, The University of North
Georgia, Gainesville, GA 30503, USA European settlement. Currently, only about 2% of this habitat
3
Present address: The Orianne Society, Indigo Snake Initiative, 414 remains, primarily in isolated fragments (Frost 2006). The
Club Drive, Hinesville, GA 31313, USA eastern indigo snake (Drymarchon couperi) is a wide-ranging

Hyslop et al.  Indigo Snake Home Range and Habitat Use 101
predator closely associated with xeric upland longleaf pine– size in indigo snakes, with males occupying larger ranges
wiregrass habitats, often referred to as sandhills, and the than females because of behavioral differences. Alternatively,
burrows of gopher tortoises (Gopherus polyphemus), which may home range size may not be directly affected by sex, but a
inhabit these upland areas. This association is especially sexually dimorphic pattern may emerge because males, on
pronounced in the northern portions of the species’ range in average, are larger (male-biased sexual size dimorphism;
south Georgia and northern Florida. Indigo snake popula- Layne and Steiner 1984; Stevenson et al. 2003, 2009), and
tions have continued to decline since federal listing in 1978 therefore have potentially greater resource needs that
(U.S. Fish and Wildlife Service [USFWS] 2008). These influence home range size (Harsted and Bunnell 1979,
declines are primarily attributed to habitat loss and Tufto et al. 1996). Patterns and scale of movement and
degradation caused by development, fire exclusion, agricul- habitat use within a landscape may be correlated with
ture, and conversion of native longleaf pine habitats to arrangement of necessary resources (Gibbons and Semlitsch
commercial plantations of off-site pine species (USFWS 1987, Gregory et al. 1987, Macartney et al. 1988). Therefore,
1978, 2008). we also predicted that variation in spatial distribution of
Gopher tortoise burrows, along with other underground resources, influenced in part by different management and
shelters, are used by indigo snakes as protection from land use objectives across the landscape, may additionally
temperature extremes, fire, and predators, and may also be affect home range size (Tufto et al. 1996, Moyer et al. 2007).
used for foraging, shelter prior to ecdysis, and nesting Habitat use in snakes may also be driven by multiple factors
(Landers and Speake 1980, Moler 1992, Stevenson similar to those influencing home range sizes, such as prey
et al. 2003, Hyslop 2007, Hyslop et al. 2009a). The and thermal requirements (Reinert 1984). We did not expect
association with tortoise burrows is especially pronounced intraspecific separation of habitat use throughout most of the
from late fall through early spring, which includes the indigo year because of assumed similarities in resource needs
snake’s breeding season from October–February (Speake between sexes; however, female indigo snakes may be limited
et al. 1978, Diemer and Speake 1983, Hyslop et al. 2009a). to using more open microhabitats for thermoregulation
Evidence suggests that indigo snakes may spend on average while gravid in the spring (Hyslop et al. 2009a). Thus, we
approximately 76% of their time underground, regardless of predicted intersexual separation in spring habitat use because
season or sex (Hyslop et al. 2009a) and that underground of differences in reproductive condition.
shelters, especially tortoise burrows, may be a limiting factor
for indigo snakes in the northern portion of their range STUDY AREA
(Diemer and Speake 1983, Hyslop et al. 2009a). Outside of We conducted a radiotelemetry study of indigo snakes on
the breeding season, and during their period of greatest Fort Stewart Military Installation (hereafter Fort Stewart)
activity with respect to movements and foraging, indigo and adjacent tracts of private land in southeastern Georgia.
snakes may use a variety of other habitats, including pine The Fort Stewart study site spanned approximately 8,000 ha
flatwoods, mixed pine-oak forests, bottomland forests, and of its total 111,600 ha (Stevenson et al. 2003); private lands
other freshwater wetlands (Speake et al. 1978, Landers and were contiguous with our Fort Stewart site and covered
Speake 1980). approximately 6,000 ha. Paved roads bordered 2 sides of the
We addressed information needs associated with the private lands, but no paved roads were present within the
eastern indigo snake in Georgia, including spatial and habitat study area. Both sites contained maintained and non-
requirements. Our objectives were to determine if eastern maintained unpaved roads. The study sites also supported
indigo snakes maintained definable home ranges, to estimate populations of gopher tortoises, although potential tortoise
annual and seasonal home range sizes and describe move- habitats on private lands were generally fire-suppressed, in
ments within home ranges, to examine ecological factors planted pine plantations, clear-cuts, and actively managed
associated with intraspecific home range size variation, and hay fields, with clusters of active burrows more localized. In
to quantify habitat use and seasonal variation in use. comparison, gopher tortoise burrows on Fort Stewart were
Home range size in eastern indigo snakes may vary more densely and uniformly distributed primarily through-
intersexually. In Georgia, female indigo snakes have been out uneven-aged pine forests.
shown to use more open microhabitats than males, especially Fort Stewart and private land study sites contained intact,
on sandhill habitats with tortoise burrows during spring longleaf pine–turkey oak (Quercus laevis)–wiregrass habitat,
(Hyslop et al. 2009a), thus resulting in intersexual differences young (5–15 yr) to medium (16–40 yr) age slash (Pinus
in home range size arising from smaller, less frequent elliottii) and loblolly (Pinus taeda) pine plantations, and
movements from females in the spring. This pattern has also recently restored (i.e., formerly in slash or loblolly pine
been shown for other snake species, and is likely attributable plantations that were clearcut) longleaf pine–wiregrass
to differences in reproductive condition and related habitats. Recent (1990–present) habitat management and
thermoregulation needs (such as increased temperatures) restoration practices at the Fort Stewart sites have promoted
when gravid (Reinert 1993, Blouin-Demers and or enhanced open-canopied longleaf pine habitats (i.e.,
Weatherhead 2001). In addition, males may also move prescribed burning from Aug to Oct, selective thinning,
greater distances when searching for mates, a behavior found longleaf pine planting, wiregrass seeding, and control of
in several other snake species (Gregory et al. 1987). xerophytic oaks via herbicide application). Other actions
Therefore, we would predict that sex influences home range included discontinuing the removal of remnant pine stumps,

102 The Journal of Wildlife Management  78(1)

which are known shelters for indigo snakes (Hyslop tracking frequency caused by seasonal variations in snake
et al. 2009a); limiting winter timber harvests on xeric activity and movement (Hemson et al. 2005).
uplands supporting tortoise burrow to avoid possible burrow We calculated annual home ranges from 15 December to
collapse at a time when snakes are more likely to be resident; 14 December 2002 to 2003 and 2003 to 2004. We estimated
and retaining debris piles (windrows) following timber seasonal MCP home ranges for winter (15 Dec–14 Mar),
harvest and site preparation, which indigo snakes have also spring (15 Mar–14 Jun), summer (15 Jun–14 Sep), and fall
been shown to use (Hyslop et al. 2009a). (15 Sep–14 Dec). We excluded individuals without complete
Management activities on non-cultivated areas of the seasons of data from that season’s analysis. We conducted
private lands included some of the same methods as Fort analyses with the Animal Movements Extension (Hooge and
Stewart; however, some sandhills on private lands had long Eichenlaub 1997) to ArcView GIS (Environmental Systems
histories of fire exclusion (>25 yr), resulting in hardwood Research Institute, Inc., Redlands, CA) and used repeated
encroachment (e.g., turkey oak), increased midstory and measures analysis of variance (ANOVA) for seasonal home
canopy cover, and reduced native ground cover. Xeric range analysis (PROC GLM; SAS Institute, Inc. 2005). We
sandhills at both sites were intergraded with mesic pine retained the individual as the sampling unit in all analyses.
flatwoods, mixed pine–oak forests, bottomland hardwood We used repeated measures linear regression on 12
habitats, and other wetlands (including isolated depressional candidate models created from a priori hypotheses to
wetlands, blackwater creek swamps, seepage-influenced bay examine biological and ecological correlates of intraspecific
swamps, and impoundments; Wharton 1978). variation of home range size (PROC MIXED; SAS
Institute, Inc. 2005). We compared models using Akaike’s
METHODS Information Criterion (AIC; Akaike 1973, Burnham and
Anderson 2002) corrected for small sample sizes (AICc;
Field Methods Hurvich and Tsai 1989). We also used AICc to select the
We captured 32 snakes, 31 by hand and 1 individual in a large covariance structure for the data from an a priori selection of
snake trap (Hyslop et al. 2009c) on upland sandhill habitats potential structures. Model averaging may be problematic
with gopher tortoise burrows (Stevenson et al. 2003) during with repeated measures designs (Reiman et al. 2006);
late fall to early spring of 2002–2004. We implanted 20 therefore, we report Akaike weights and parameter estimates
snakes (7 F, 13 M) with radiotransmitters between for model parameters included in 90% model confidence sets.
December 2002 and April 2003 and 12 additional snakes Model parameters included sex, snout-vent length (size),
(6 F, 6 M) from October 2003 to March 2004. For details of interaction of sex and size, overwintering location (Fort
surgical and care procedures see Hyslop et al. (2009b). We Stewart or private lands), and number of radio locations. We
located snakes 2–3 times per week by foot and vehicle using standardized size by sex using residuals of size versus sex
homing techniques (Mech 1983), with accuracy <1 m if the regression as an additional covariate in our models (size
animal was underground or immobile (e.g., basking; standardized). In addition to setting a minimum tracking
behavior noted if individual was observed). We staggered duration for inclusion of individuals into annual and seasonal
periods randomly during the day that we tracked each home range and bootstrap analyses, we also incorporated the
individual. Animal Care and Use Committees of The number of locations collected per individual to evaluate if
University of Georgia IACUC (A2002-10111-0) and home ranges were maintained.
USGS Patuxent Wildlife Research Center approved study
procedures. Patterns of Movement
To calculate frequency of movement, we divided the
Home Range 24 months of radiotelemetry data into 14-day periods and
To examine area used by the snakes, we used 100% minimum calculated proportion of days moved compared with number
convex polygons (MCP) to estimate home ranges of days radiotracked within that 14-day period. This method
(Mohr 1947, Southwood 1966) and kernel density (KD) standardized tracking effort across seasons and individuals.
analysis to estimate utilization distributions (Worton 1989). We deleted records for snakes with only 1 novel location
To address possible sample size bias associated with MCPs during a 14-day period (31 deleted locations, 18 individuals).
(e.g., Arthur and Schwartz 1999), we used bootstrap analysis To generate a minimum daily movement index, we
(500 iterations) to examine sample size to home range area calculated straight-line distances between successive loca-
relationships and included home ranges in analysis only if tions (Animal Movements Extension; Hooge and
incremental area curves visually reached an asymptote. We Eichenlaub 1997). We based the calculations on the number
calculated KD at 95% isopleths (home range) and at 50% of days in each season that individual snakes were located; we
isopleths (core activity areas; Samuel et al. 1985) using the again averaged movement for each individual during 14-day
fixed kernel method with a least squares cross-validation periods to standardize data. For movement analyses, we used
smoothing parameter (Worton 1989, Seaman and repeated measures ANOVA and retained the individual as
Powell 1996, Gitzen and Millspaugh 2003). For calculation the sampling unit (SAS Institute, Inc. 2005).
of KD home ranges, we retained only novel radiolocations
and removed consecutive repeated locations (individual in Habitat Association Analyses
same underground shelter or within approx. 2 m of previous Conclusions regarding habitat use may vary with spatial scale
location if on the surface) to alleviate potential bias in examined; therefore, we analyzed habitat use hierarchically

Hyslop et al.  Indigo Snake Home Range and Habitat Use 103
(Johnson 1980). We examined the snake’s position within its clearcut, sandhill, pine plantation, and miscellaneous uplands
home range by comparing habitat at telemetry locations to as described in the in-field habitat category explanations. We
habitat within MCP home ranges (site selection) and used repeated measures ANOVA with arcsine transformed
compared proportional habitat composition within home data and a Tukey-Kramer multiple comparison procedure for
ranges to proportion of habitats available at the site (home examining seasonal differences in habitat use between males
range selection; Johnson 1980). We used compositional and females (PROC GLM; SAS Institute, Inc. 2005).
analysis (Aitchison 1986, Aebischer et al. 1993) for habitat
use comparisons (Bycomp Version 1.0; Ott and Hovey 1997; RESULTS
SAS Institute, Inc. 2005). We categorized Gap Analysis
Program land cover data (GAP; Kramer et al. 2003) into 7 We collected 4,993 telemetry locations for 32 snakes from
major land cover types: 1) roads and urban areas (road- January 2003 to December 2004 (24 months). Individuals
urban); 2) open water, forested bottomland wetlands, were tracked 89–711 days (x ¼ 420 days, 39–254 locations
depressional wetlands, bay swamps, and non-forested wet- per individual). Male snout-vent length averaged 158 cm
lands (wetlands); 3) agricultural and other fields (field); 4) (range ¼ 120–191) and mass at capture averaged 2.2 kg
clearcuts and other habitats with sparse canopy cover, (range ¼ 0.7–4.3). Female snout-vent length averaged
including regenerating forests generally <10 years in age 138 cm (range ¼ 110–156) with an average mass of 1.5 kg
with low canopy cover (sparse); 5) forests with >75% (range ¼ 0.6–2.3).
deciduous trees generally >10 years in age (deciduous); 6) Home Range
forests with >75% evergreen trees, including managed pine Bootstrap analysis of MCP annual and seasonal home ranges
plantations generally >10 years in age (evergreen); and 7) yielded area curves that reached asymptote, suggesting
pine-hardwood mixed forest, including shrub-scrub habitats sufficient locations were collected for the home range area to
(mixed). stabilize. We included 18 snakes in 2003 (11 M, 7 F) and 20
At each snake radiolocation, we also measured habitat snakes in 2004 (13 M, 7 F) in annual home range calculations
characteristics in the field and defined in-field habitat (13 snakes from 2003 also included in 2004). Annual home
categories based on hydrology, vegetation, management, and ranges (MCP) averaged 378 ha (95% CI ¼ 185–571) in 2003
presence of gopher tortoise burrows. We included this and 340 ha (95% CI ¼ 184–496) in 2004, with individual
additional habitat use analysis primarily to distinguish home ranges from 33–354 ha for females and from 140–
sandhill habitats from evergreen, mixed, and sparse GAP 1,528 ha for males. Males, on average, occupied annual home
categories, each of which contained some sandhill habitat. ranges approximately 4.5 times larger than females in 2003
In-field habitat categories included 1) xeric uplands with and approximately 6.6 times larger than females in 2004 for
longleaf pine and/or xerophytic oak canopy and gopher MCP and KD (Table 1). Kernel density analysis at 50%
tortoise burrows (sandhill); 2) harvested pine flatwoods with isopleths yielded 1–5 distinct regions of core habitat use for
windrows, low to no canopy cover, bedding for loblolly pine each snake (x ¼ 1.7), with no differences between sexes or
planting, embedded isolated wetlands, without tortoise years. Males, on average, also occupied larger individual core
burrows (clearcut); 3) old-field, hay fields, and food plots, activity areas than females; although core areas averaged 12%
some areas with tortoise burrows (field); 4) pine plantations of 95% KD home ranges regardless of sex. All annual home
of various ages, may or may not contain tortoise burrows ranges in our sample overlapped with 6 other home ranges,
(pine plantation); 5) areas of mixed pine-hardwood canopy with no evidence or inter- or intraspecific avoidance.
composition, occasional tortoise burrows (miscellaneous Global models for MCP, 95% KD, and 50% KD annual
uplands); and 6) low-lying seasonally flooded areas with home ranges confirmed adequate fit (likelihood ratio test:
no tortoise burrows (wetlands). P < 0.05). Residual normality plots of natural log-trans-
The in-field habitat categories field and wetland were formed data supported normality of transformed data in
similar to the GAP land cover categories of the same name. annual home range estimates. Analyses of global models
No in-field category corresponded with the road-urban GAP suggested autoregressive covariance structure was the
category because we did not record any snake locations in this most appropriate. The 90% confidence set of models for
habitat type. We separated the evergreen, mixed, deciduous, estimating annual MCP home ranges contained 2 of 12
and sparse GAP categories into the in-field categories candidate models (Table 2). The model with the most
Table 1. Home range and activity center areas for male (M) and female (F) eastern indigo snakes radiotracked 2003–2004, Georgia. Values (ha) are averages
with 95% confidence intervals (CI) for 100% minimum convex polygon (MCP) home ranges, 95% kernel density (KD) utilization distributions, 50% KD
activity centers, and average number of distinct 50% KD core areas. Of the snakes tracked in 2003 with complete years of data (n ¼ 18), 13 of those were also
tracked in 2004 (11 M, 2 F).
100% MCP 95% KD 50% KD
Year Sex n x 95% CI x 95% CI x 95% CI Average number of core areas
2003 M 11 538 264–812 792 390–1,194 120 21–219 1.7
F 7 126 45–207 173 59–587 25 6–44 1.6
2004 M 13 482 279–685 552 242–862 76 42–110 1.6
F 7 77 57–97 79 54–104 10 7–13 1.8

104 The Journal of Wildlife Management  78(1)

Table 2. Candidate models for annual 100% minimum convex polygon (MCP) home ranges for radiotracked eastern indigo snakes (n ¼ 32), 2003–2004,
Georgia. Models are listed in order of corrected Akaike’s Information Criterion (AICc), with number of parameters (K), difference in AICc (DAICc), model
likelihood, and Akaike weights (v) for the set of candidate models (i).
Modela K AICc ~AICc Model likelihood vi
Sex, size, site 6 114.60 0.00 1.00 0.824
Sex, size 5 118.26 3.66 0.16 0.132
Sex, size, sex  size 6 121.40 6.80 0.03 0.027
Sex, size, site, locations, sex  size 8 122.40 7.80 0.02 0.017
Sex, site 5 136.46 21.86 0.00 0.000
Sex 4 143.82 29.22 0.00 0.000
Size, site 5 144.06 29.46 0.00 0.000
Size 4 148.62 34.02 0.00 0.000
Site 4 175.02 60.42 0.00 0.000
Size (standardized), site 5 173.66 59.06 0.00 0.000
Locations 4 184.22 69.62 0.00 0.000
Size (standardized) 4 181.82 67.22 0.00 0.000

a
Model parameters: sex (being female), size (snout-vent length), site (overwintering location on Fort Stewart vs. private lands), locations (number of telemetry
locations), and size (standardized; snout-vent length standardized by sex).

support included sex, size, and overwintering site (v ¼ 0.824) Several large-ranging males (n ¼ 3) traveled 5–8 km linear
and was 6.2 times more likely than the next best distance from winter to summer locations. One of these
approximating model, which contained sex and size males used the same travel corridor (approx. 2 km) for 2
(v ¼ 0.132). Parameter estimates were similar between the consecutive years, despite about 75% of the corridor having
2 models, suggesting smaller home ranges for females, for been clearcut in the intervening winter. Although we
smaller individuals, and for individuals overwintering on recorded 6 individuals <100 m from paved roads, no snake
Fort Stewart (Table 3). Results were similar for 95% KD locations were outside boundaries created by paved roads.
annual home ranges and 50% KD activity centers. Sex ranked During the study, however, 2 indigo snakes not included in
as the most influential factor modeled, according to Akaike the telemetry study were found dead on these roads.
importance weights, for all annual home range estimates; size Radiotracked snakes, however, crossed unpaved roads and
ranked second and site ranked third (Table 4). trails regularly on Fort Stewart and private lands.
Seasonal home range size varied annually (F7,109 ¼ 49.8, Average daily distance moved (Fig. 2), averaged biweekly,
P < 0.001) and by sex (F1,30 ¼ 34.8, P < 0.001), with no varied by sex (F1,30 ¼ 14.7, P < 0.001) and season
interaction (F7,109 ¼ 0.24, P ¼ 0.97; repeated measures (F3,84 ¼ 79.3, P < 0.001) with no interaction (F3,84 ¼ 1.8,
ANOVA). In the winter, home ranges were smallest and P ¼ 0.16). Females had smaller daily movement distances
most similar in size for males and females (Fig. 1). than males, regardless of season (Fig. 2). Winter movements
were smaller than other seasons, with no difference between
Patterns of Movement sexes (t84 ¼ 0.1, P ¼ 0.91). Average frequency of biweekly
All snakes tracked 9 months returned to sandhills used the movement also varied by sex (F1,30 ¼ 4.6, P ¼ 0.04) and
previous fall and winter. Individual degrees of fidelity to season (F3,81 ¼ 65.8, P < 0.001) with an interaction effect
specific shelters used by each snake varied; however, all (F3,81 ¼ 5.9, P  0.001). Males moved more often than
snakes tracked in fall and winter 2003–2004 and 2004–2005 females in all seasons (Fig. 2). Least squares estimates of
returned to a minimum of 4 different underground shelters differences in movement frequency indicated similar
that they had specifically used the preceding fall and winter. movement patterns in fall and spring for females

Table 3. Estimates of fixed and random effects for the 90% confidence set of models examining minimum convex polygon (MCP) home ranges of
radiotracked eastern indigo snakes (n ¼ 32), 2003–2004, Georgia. Data suggest a negative effect of being female and a positive effect of body size on home
range size.
Modela Effect Parameter Estimate Lower 95% CL Upper 95% CL
Sex, size, site Fixed Sex 0.985 1.423 0.547
Size 0.021 0.009 0.033
Site 0.382 0.794 0.029
Random Intercept 2.786 0.948 4.624
Residual 0.245 0.159 0.938
Year (repeated) 0.610 0.282 0.427

Sex, size Fixed Sex 1.050 1.510 0.591

Size 0.024 0.011 0.036
Random Intercept 2.220 0.393 4.048
Residual 0.276 0.179 0.482
Year (repeated) 0.657 0.371 0.944

a
Model parameters: sex (being female), size (snout-vent length), and site (overwintering location on Fort Stewart vs. private lands).

Hyslop et al.  Indigo Snake Home Range and Habitat Use 105
Table 4. Influence of sex and size on variation of intraspecific home range most to least selected included evergreen, wetland, mixed,
size in eastern indigo snakes radiotracked (n ¼ 32) in Georgia, 2003–2004.
Data are Akaike importance weights for model parameters from annual
sparse, field, deciduous, and roads. Selection of the top 4 land
minimum convex polygon (MCP) home ranges, 95% kernel density (KD) cover types was significantly different from the last 3 habitats
home ranges, and 50% KD core activity areas. (Table 5).
Importance weights Habitat use varied seasonally for in-field habitat types
(Fig. 3). In winter, snakes restricted their habitat use
Candidate Annual Annual Annual
Parametersa models MCP 95% KD 50% KD primarily to sandhills with gopher tortoise burrows (sand-
Sex 6 1.00 1.00 1.00
hill). About 67% of all locations in winter were in sandhills
Size 6 0.99 0.99 0.98 and 22% occurred in young, upland pine plantations with
Site 5 0.84 0.71 0.39 gopher tortoise burrows (pine plantation). Snakes exhibited
Size  sex 2 0.04 0.19 0.12 less concentration in use of sandhill habitats in spring,
Locations 2 0.02 — —
Size (standardized) 2 0.00 0.00 0.00
summer, and fall than during winter; however, snakes
continued to use sandhill habitats throughout the warm
a
Model parameters: sex (being female), size (snout-vent length), site months ranging from 35% to 58% of locations depending on
(overwintering location on Fort Stewart vs. private lands), locations the season (Fig. 3). Use of wetlands in summer (x ¼ 30%,
(number of telemetry locations), and size (standardized; snout-vent 95% CI ¼ 0.23–0.36) was greater than recorded in any other
length standardized by sex).
season and was concentrated primarily in bottomland
hardwood areas between sandhills and other upland
(t1,81 ¼ 0.8, P ¼ 0.44), but not for males (t1,81 ¼ 3.3, habitat types.
P ¼ 0.003). We recorded more frequent movements in Use of in-field habitat categories also varied within
summer compared to other seasons. Snakes moved least each season (winter F6,189 ¼ 30.0, P < 0.001; spring
often in winter with no difference between sexes (t1,81 ¼ 0.5, F6,196 ¼ 45.22, P < 0.001; summer F6,182 ¼ 20.6, P < 0.001;
P ¼ 0.60; Fig. 2). fall F6,168 ¼ 28.1, P < 0.001) with no differences between
male and female habitat use detected, but with a sex  season
Habitat Use interaction effect in spring (F6 ¼ 3.09, P ¼ 0.006). Tukey–
Average use of GAP land cover types across the study site Kramer post hoc tests of habitat use in spring indicated that
(average proportion within home ranges [proportion females used sandhills more than males (P ¼ 0.04), with no
available across study sites]) were 0.01 (0.02) road-urban, other seasonal differences in habitat use between sexes.
0.26 (0.24) wetlands, 0.05 (0.07) fields, 0.08 (0.07) sparse,
0.02 (0.03) deciduous forest, 0.49 (0.51) evergreen, and 0.09 DISCUSSION
(0.06) mixed. Compositional analysis indicated nonrandom
use of habitat at both levels of selection examined (home Home Range
range selection: l ¼ 0.212, P < 0.001; site selection: Results of bootstrap analyses and modeling suggest our
l ¼ 0.324, P ¼ 0.01). Use of habitats in home range selection telemetry efforts were sufficient to describe home ranges
ranked in descending order was wetland, evergreen, mixed, and indicated that individuals maintained definable annual
field, sparse, road-urban, and deciduous. Selection differed home ranges. The male indigo snake home ranges (140–
between the 5 most selected habitats and the 2 least selected 1,528 ha) reported herein may represent the largest reported
habitats. Habitat use rankings for site selection analysis, from for an indigenous terrestrial North American snake species

500 13 Male
Female
400 13
100% MCP (ha)

13 13
300
14
10
200

7
100 7 9
7 9 9
5 3 17 11
0
Winter Spring Summer Fall Winter Spring Summer Fall
2003 2004

Figure 1. Seasonal and sex differences in 100% minimum convex polygon (MCP) home ranges (x, 95% CI) for male (gray bars) and female (white bars) eastern
indigo snakes radiotracked for complete seasons, 2003–2004, Georgia. Values represent sample size for each estimate. Males, outside of winter, had seasonal
home ranges approximately 4.5 times larger than females.

106 The Journal of Wildlife Management  78(1)

 Females Males species, have reported MCP home ranges of <1 ha to 183 ha
180 (n ¼ 4; Dodd and Barichivich 2007). The eastern diamond-
Average movement distance (m)

160 back rattlesnake (Crotalus adamanteus), another large species,

140 has home ranges reported between 28 ha and >80 ha (see
120 Waldron et al. 2008 for review). Home ranges (MCP) of
100 male timber rattlesnakes (Crotalus horridus) are reported
80 averaging around 112 ha (Macartney et al. 1988, Reinert and
60 Zappalorti 1988).
40
Previous studies of eastern indigo snakes in northern
20
portions of the range reported individual home range sizes of
0
5–230 ha (Speake et al. 1978, Smith 1987, Speake 1993);
J F M A M J J A S O N D
0.9
however, these results included data from translocated and
0.8
captive-reared individuals, making comparisons to our
Average movement frequency

0.7
results problematic. In central and coastal peninsular Florida,
0.6 cumulative home ranges (100% MCP), for periods between
0.5 10 months and over 2 years, were 39–583 ha for radiotracked
0.4 male indigo snakes (x ¼ 202, n ¼ 23) and 13–313 ha for
0.3 females (x ¼ 76, n ¼ 21; Breininger et al. 2011). In northern
0.2 peninsular Florida, annual home ranges for males (n ¼ 4)
0.1 averaged 141 ha (32–281 ha; Moler 1985) and 185 ha for 1
0 male tracked for 322 days (Dodd and Barichivich 2007).
J F M A M J J A S O N D
Month
Models evaluating potential sources of variation for home
range area suggested a strong relationship with sex (females
Figure 2. Seasonal and sex differences in daily movement distance (top) and have smaller home ranges on average) and a slight positive
average movement frequency (bottom) consistent with seasonal home range effect of increasing body size (larger snakes have larger home
patterns. We calculated values biweekly for radiotracked male (black) and
female (gray) eastern indigo snakes (n ¼ 32 total), 2003–2004, Georgia.
ranges on average). Although indigo snakes are male-biased
Individual animals were retained as the sampling unit for calculations. sexually size dimorphic, the snakes in our sample exhibited
considerable overlap in body size between larger adult
females and smaller adult males. Reproductive condition and
(Macartney et al. 1988, Reinert and Zappalorti 1988, Dodd associated behaviors may account for some differences in
and Barichivich 2007, Kapfer et al. 2008, Waldron home range size between sexes, especially in the spring when
et al. 2008). In a review of the genus Pituophis (pine and females remained on sandhill habitats where they over-
bullsnakes), Kapfer et al. (2008) reported male home range wintered, whereas males dispersed into a variety of habitats,
sizes averaged 79 ha (MCP). Eastern coachwhips (Mastico- including wetlands. Size standardized by sex had no
phis flagellum flagellum), another large North American snake predictive power in our models, further suggesting that

Table 5. Differential use of habitats compared to availability within the study site and within individual home ranges for radiotracked eastern indigo snakes
(n ¼ 27), 2003–2004, Georgia. Data present the log-ratio matrix of differences in use between GAP land cover types calculated as the log of the ratio between
the relative use of habitat types for home range selection (comparison of habitat within individual home ranges to habitat available at the study site) and site
selection (comparison of habitat at radiolocations to habitat available within individual home ranges). Positive values indicate the habitat in the column was
used relatively more than habitat in the row; negative values indicate less use. An asterisk denotes deviation from random at P < 0.05. Rank 1 represents the
most used relative to available habitat to the study animals when comparing relative use to availability; rank 7 represents the least.
Road-urban Wetland Field Sparse Deciduous Evergreen
Rank x SE x SE x SE x SE x SE x SE
Home range selection
Road-urban 6
Wetland 1 1.13 0.30
Field 3 0.01 0.46 0.71 0.45
Sparse 3 0.64 0.43 0.72 0.40 0.66 0.51
Deciduous 7 0.14 0.61 1.10 0.38 0.74 0.51 0.38 0.47
Evergreen 2 1.17 0.26 0.05 0.14 0.92 0.47 0.67 0.34 1.05 0.44
Mixed 3 0.49 0.56 0.51 0.32 0.16 0.43 0.22 0.48 0.59 0.26 0.46 0.38

Site selection
Road-urban 7
Wetland 2 1.51 0.38
Field 5 0.45 0.69 1.22 0.51
Sparse 4 1.03 0.60 0.31 0.51 1.06 0.78
Deciduous 6 0.31 0.72 1.25 0.49 0.05 0.78 0.94 0.71
Evergreen 1 1.47 0.41 0.04 0.17 1.05 0.52 0.35 0.51 1.29 0.48
Mixed 3 1.12 0.59 0.02 0.43 0.48 0.75 0.29 0.56 1.23 0.67 0.06 0.40

Hyslop et al.  Indigo Snake Home Range and Habitat Use 107
 0.8 Winter
Spring
0.7
Summer

0.6 Fall

0.5
.
Proportion

0.4

0.3

0.2

0.1

0
Sandhill Plantation Wetland Upland Clearcut Field

Figure 3. Proportionally high use of sandhill habitats annually and wetland habitats in summer (x, 95% CI) for eastern indigo snakes radiotracked in 2003–
2004, Georgia (nwinter ¼ 31, nspring ¼ 32, nsummer ¼ 28, nfall ¼ 28). Habitat types recorded at radiolocations: sandhill (pine-oak xeric uplands with longleaf pine
canopy and gopher tortoise burrows), pine plantation (included windrows), wetlands, miscellaneous uplands (xeric uplands with mixed canopy composition),
clearcut, and field (old-fields, maintained hay fields, food plots). Individual animals were the sampling unit for calculations.

intersexual differences, rather than differential resource individuals, at least during the non-breeding period. All
needs for larger individuals, affected home range size snake home ranges, however, overlapped those of multiple
variation between sexes. other snakes (n  6) both temporally and spatially, and the 2
Analyses also indicated an increase in home range size with largest home ranges (both males) occurred in close proximity
indigo snakes overwintering on private lands (MCP models) to and overlapped each other in both summer and winter.
or with an increasing proportion of locations on private lands Home ranges in other snake species that have been shown to
(KD models). Our results suggest a possible effect of habitat vary by sex have been attributed to differential energetic
type and land use on home range size indicating that needs and reproductive condition (Gregory et al. 1987,
although necessary resources were likely available on private Whitaker and Shine 2003). Eastern indigo snakes are active
lands, they may have existed in lower densities than those at foragers; we recorded approximately 84% of foraging
Fort Stewart. Confidence intervals for the site variable, observations (n ¼ 65 observations) in spring and summer
however, spanned 0 in all models, lending uncertainty (also see Stevenson et al. 2010 for review), when snakes
regarding its influence (Table 3). The site variable was not exhibited much larger and more frequent movements than in
ranked as high in models for 50% KD cores of activity as with cooler months. Approximately 65% of observations of
95% KD and MCP models (Table 4). Cores of activity (50% foraging or prey consumption behavior were in wetlands
KD) were primarily located around individual overwintering and 20% in sandhills. Therefore, in addition to effects of sex
areas, which indicated little influence of site on the size of and body size, availability of prey resources likely influences
cores of activity and overwintering areas. indigo snake home ranges at least in warmer months.
Consistent effects for body size on home range size have Indigo snakes maintained their smallest home ranges in
not been established inter- or intra-specifically in snakes winter, intermediate-sized in spring and fall, and largest in
(Gregory et al. 1987, Macartney et al. 1988). Large home summer. Previous radiotelemetry of indigo snakes in Georgia
range sizes such as those observed in this study, however, may reported the smallest home ranges from December to April
represent the needs of a large terrestrial predator, which on (x ¼ 4.8 ha, n ¼ 8), intermediate-sized from May to July
average, requires more food and area to forage than a smaller (x ¼ 42.9 ha, n ¼ 9), and largest from August to November
species (McNab 1963) or potentially better quality habitat (x ¼ 97.4 ha, n ¼ 4; Speake et al. 1978). The annual trend is
with greater density of prey available. If these large home similar in both studies, although direct comparisons are not
ranges were an artifact of snakes seeking new overwintering possible because some translocated snakes were used and
and breeding areas (i.e., emigration), we would expect that a seasons were designated differently in the previous study.
proportion of snakes would not return to the overwintering
area used the previous year. All snakes tracked in our study Patterns of Movement
returned to the same area used the previous winter, a pattern Patterns of movement were consistent with home range data,
observed during 3 winters of telemetry. A mark–recapture reflecting seasonality in movements and differences between
study at Fort Stewart, however, documented occasional use sexes. Many snake species exhibit larger and more frequent
of multiple overwintering sites between years by at least 4 of movements during the breeding season (e.g., Gibbons and
63 marked male snakes (Stevenson et al. 2009). Dorcas 2004); however, snakes in this study exhibited the
If intraspecific competition influenced home range sizes in opposite pattern. In our study population, breeding occurs
this study, we would expect to see some avoidance between when adult snakes congregate on xeric sandhills during or

108 The Journal of Wildlife Management  78(1)

just prior to the coldest months of the year when movements were mesic pine flatwoods prior to timber harvesting and
are greatly reduced compared to other seasons (Fig. 3). contained bedding for new plantings. Approximately 65% of
Seasonally, we found fewer differences in movement observations of foraging behavior or prey consumption were
frequency between males and females outside of spring in wetland habitats. During summer, males and females
than for movement distances, suggesting similar vagility depended less on sandhill habitats, but continued to use these
throughout much of the year, but at a reduced spatial scale for habitats and tortoise burrows for shelter during ecdysis and
females. often following foraging (Hyslop et al. 2009a). In fall, habitat
Indigo snakes radiotracked in this study followed 2 general use was transitional between summer and winter use, as
movement patterns. All females and approximately half the snakes began to return to overwintering sandhills.
males maintained associations with their overwintering Ecosystem approaches to management often focus on a few
sandhill sites throughout the year, despite greater use of non- key species that may serve as indicators of ecosystem integrity
sandhill habitats in warmer seasons. Males with the largest (Lambeck 1997, Roberge and Angelstam 2004). Wildlife
home ranges in this study made directional movements of management for nongame species on many public lands in
between 1,500 m and 7,500 m in late spring and did not the southeastern United States with longleaf pine habitats
return to their respective overwintering sandhills until mid- has often focused on red-cockaded woodpecker (Picoides
fall (n ¼ 6). We considered both of these movement patterns borealis) and gopher tortoise populations (Fort Stewart 2001,
home ranges because of return to overwintering sandhills and USFWS 2008). These activities often include maintaining a
asymptotic results from bootstrapping analyses (Burt 1943). low basal area of trees, sparse canopy cover, low midstory
cover, native groundcover vegetation, and prescribed fire
Habitat Use (Brockway et al. 2005, FFWCC 2007). Although these
Our data from southeastern Georgia indicate that eastern management strategies are likely to benefit indigo snakes, if
indigo snakes exhibit specific affinities for open-canopied the habitat complementation needs (Dunning et al. 1992,
sandhill habitats supporting gopher tortoise populations in Pope et al. 2000) of indigo snakes are not considered (such as
the winter and for lowlands including expansive wetlands in adjacent wetlands), then current management strategies and
the warmer months. Habitat use analyses conducted on GAP land acquisition plans may be inadequate for indigos and
land cover categories indicated that wetlands, sparse, mixed, other snake species (Steen et al. 2012).
and evergreen forests were used at greater rates than expected
relative to their availability at both spatial scales examined. MANAGEMENT IMPLICATIONS
Assessment of sparse areas used by snakes revealed some use Habitat loss, fragmentation, and degradation remain primary
of clearcut sites (e.g., former mesic pine flatwoods) for males threats to indigo snake populations (USFWS 2008). This
in warmer months, but most areas were predominantly study adds to our understanding of the space use and
comprised of young longleaf pine plantations supporting potential habitat requirements of the species. First, indigo
gopher tortoise populations, especially in winter. snakes may require more land than previously recommended.
Of the habitat types recorded in the field, sandhills were Although land area requirements were not directly
used more than any other habitat for both males and females addressed, a recent review of the status of the species
in all seasons, except for summer when wetland and sandhill (USFWS 2008) noted a reserve size of 4,046 ha as likely
use was comparable. Winter included breeding activities on beneficial for indigo snakes, whereas others have recom-
upland habitats, extended periods of inactivity, and use of mended multiple 1,000-ha tracts of land for conservation of
tortoise burrows during cold temperatures (Hyslop the species in Florida (Moler 1992). The collective extent of
et al. 2009a). During spring, males began dispersing from the radiolocations for the 32 snakes in this study spanned an
sandhills to surrounding habitats, including wetlands, area of 8,000 ha (MCP of all locations) to 14,000 ha
clearcuts, and other upland habitats. Females remained on (rectangular area). Second, longleaf pine-wiregrass habitats
sandhill habitats until late spring and early summer when exist as a natural mosaic of upland and lowland areas that
oviposition is completed (Speake et al. 1987). In spring 2004, require frequent, low to moderate intensity fire to maintain
ultrasound and/or radiographs on 9 of 10 females in the study biological diversity (Mushinsky and McCoy 1985, Van Lear
at that time showed signs of initial egg formation (Hyslop et al. 2005). Because development and agriculture has
et al. 2009b), supporting our prediction that greater use of open eliminated natural habitats between sandhills, islands of
habitats by females was associated with thermoregulatory remnant habitat within an unsuitable matrix may have little
requirements during gestation. Similar patterns of movement to no connectivity (McCoy and Mushinsky 1999). Move-
have been reported of earlier male dispersal from overwintering ment patterns and habitat use in this study support variable
locations compared to females (Parker and Brown 1980, permeability of these matrix types to indigo snake
Shine 2003); however, few studies have been conducted on movement, implicating that patch connectivity may be as
temperate species, such as the eastern indigo snake, that are important as habitat patch size for population persistence.
frequently surface-active throughout cooler seasons. Thus, habitat fragmentation, roads, and land development,
Both Fort Stewart and private land sites had extensive even at low densities, may exacerbate impacts of habitat loss
bottomland hardwood habitats that were used extensively by on indigo snakes because of their use of multiple habitats,
snakes. Other wetlands used, although to a lesser extent, large home ranges, and seasonal movements within those
included cypress domes embedded in large clearcuts, which ranges (Breininger et al. 2012). Given our results, we propose

Hyslop et al.  Indigo Snake Home Range and Habitat Use 109
that the inclusion of eastern indigo snakes into biodiversity Florida Fish and Wildlife Conservation [FFWCC]. 2007. Gopher tortoise
management and conservation in the southeastern Coastal management plan. Florida Fish and Wildlife Conservation, Tallahassee,
USA.
Plain would benefit a broad range of species (Grumbine Fort Stewart Directorate of Public Works. 2001. Integrated natural
1994, Noss 2000). resources management plan, 2001–2005. Fort Stewart and Hunter Army
Airfield, Fort Stewart, Georgia, USA.
Frost, C. 2006. History and future of the longleaf pine ecosystem. Pages 9–
ACKNOWLEDGMENTS 42 in S. Jose, E. J. Jokela, and D. L. Miller, editors. The longleaf pine
ecosystem: ecology, silviculture, and restoration. Springer, New York,
The Georgia Department of Natural Resources, Nongame New York, USA.
Wildlife and Natural Heritage Section (Section 6 funding Gibbons, J. W., V. J. Burke, J. E. Lovich, R. D. Semlitsch, T. D. Tuberville,
by U.S. Fish and Wildlife Service), and United States J. R. Bodie, J. L. Greene, P. H. Niewiarowski, H. H. Whiteman, D. E.
Scott, J. H. K. Pechmann, C. R. Harrison, S. H. Bennett, J. D. Krenz,
Geological Survey Patuxent Wildlife Research Center M. S. Mills, K. A. Buhlmann, J. R. Lee, R. A. Seigel, A. D. Tucker, T. M.
funded this research. We thank the wildlife biologists and Mills, T. Lamb, M. E. Dorcas, J. D. Congdon, M. H. Smith, D. H.
land managers of Fort Stewart for their considerable Nelson, M. B. Dietsch, H. G. Hanlin, J. A. Ott, and D. J. Karapatakis.
assistance throughout this project, particularly T. Beaty, 1997. Perceptions of species abundance, distribution, and diversity: lessons
from four decades of sampling on a government-managed reserve.
and L. Carlile. We gratefully acknowledge J. Jensen and M. Environmental Management 21:259–268.
Harris for support of this project, J. Caligiure for base access Gibbons, J. W., and M. E. Dorcas. 2004. North American watersnakes: a
assistance, and J. Thrift. T. Norton, Wildlife Conservation natural history. University of Oklahoma Press, Norman, USA.
Society, and St. Catherines Island Foundation provided Gibbons, J. W., and R. D. Semlitsch. 1987. Activity patterns. Pages 396–421
in R. A. Seigel, J. T. Collins, and S. S. Novak, editors. Snakes: ecology and
invaluable services and assistance with implantation proce- evolutionary biology. MacMillan, New York, New York, USA.
dures. We also thank 2 anonymous reviewers, J. W. Gibbons, Gitzen, R. A., and J. J. Millspaugh. 2003. Comparison of least-squares cross-
C. Hass, and D. Steen for comments and suggestions that validation bandwidth options for kernel home-range estimation. Wildlife
improved the manuscript. Society Bulletin 31:823–831.
Gregory, P. T., J. M. MaCartney, and K. W. Larsen. 1987. Spatial patterns
and movements. Pages 366–395 in R. A. Seigel, J. T. Collins, and S. S.
LITERATURE CITED Novark, editors. Snakes: ecology and evolutionary biology. MacMillan,
Aebischer, N. J., P. A. Robertson, and R. E. Kenward. 1993. Compositional New York, New York, USA.
analysis of habitat use from animal radio-tracking data. Ecology 74:1313– Grumbine, R. E. 1994. What is ecosystem management? Conservation
1325. Biology 8:27–38.
Aitchison, J. 1986. The statistical analysis of compositional data. Chapman Harsted, A. S., and F. A. Bunnell. 1979. Home range and body weigh—a
& Hall, London, United Kingdom. reevaluation. Ecology 60:389–402.
Akaike, H. 1973. Information theory and an extension of the maximum Hemson, G., P. Johnson, A. South, R. Kenward, R. Ripley, and D.
likelihood principle. Pages 267–281 in B. N. Petrov and F. Csaki, editors. Macdonald. 2005. Are kernels the mustard? Data from global positioning
Second International Symposium of Information Theory. Akademiai system (GPS) collars suggests problems for kernel home-range analyses
Kiado, Budapest, Hungary. with least-squares cross-validation. Journal of Animal Ecology 74:455–
Arthur, S. M., and C. C. Schwartz. 1999. Effects of sample size on accuracy 463.
and precision of brown bear home range models. Ursus 11:139–148. Hooge, P. N., and W. M. Eichenlaub. 1997. Animal movement extension to
Blouin-Demers, G., and P. J. Weatherhead. 2001. Thermal ecology of black ArcView. U.S. Geological Survey, Alaska Biological Science Center,
rat snakes (Elaphe obsoleta) in a thermally challenging environment. Anchorage, USA.
Ecology 82:3025–3043. Hurvich, C. M., and C. L. Tsai. 1989. Regression and time-series model
Breininger, D. R., M. R. Bolt, M. L. Legare, J. H. Drese, and E. D. Stolen. selection in small samples. Biometrika 76:297–307.
2011. Factors influencing home-range sizes of eastern indigo snakes in Hyslop, N. L. 2007. Movements, habitat use, and survival of the threatened
central Florida. Journal of Herpetology 45:484–490. eastern indigo snake (Drymarchon couperi) in Georgia. Dissertation, The
Breininger, D. R., M. J. Mazerolle, M. R. Bolt, M. L. Legare, J. H. Drese, University of Georgia, Athens, USA.
and J. E. Hines. 2012. Habitat fragmentation effects on annual survival of Hyslop, N. L., R. J. Cooper, and J. M. Meyers. 2009a. Seasonal shifts in
the federally protected eastern indigo snake. Animal Conservation 15: shelter and microhabitat use of the threatened eastern indigo snake
361–368. (Drymarchon couperi) in Georgia. Copeia 2009:460–466.
Brockway, D. G., K. W. Outcalt, D. J. Tomczak, and E. E. Johnson. 2005. Hyslop, N. L., J. M. Meyers, R. J. Cooper, and T. M. Norton. 2009b.
Restoring longleaf pine forest ecosystems in the southern US. Pages 501– Survival of radio-implanted eastern indigo snakes (Drymarchon couperi) in
519 in J. A. Stanturf and P. Madsen, editors. Restoration of boreal and relation to body size and sex. Herpetologica 65:199–206.
temperate forests. CRC Press, Boca Raton, Florida, USA. Hyslop, N. L., J. M. Meyers, R. J. Cooper, and D. J. Stevenson. 2009c.
Burnham, K. P., and D. R. Anderson. 2002. Model selection and Indigo snake capture methods: effectiveness of two survey techniques for
multimodel inference: a practical information-theoretical approach. Drymarchon couperi in Georgia. Florida Scientist 72:93–100.
Second edition. Springer-Verlag, New York, New York, USA. Johnson, D. H. 1980. The comparison of usage and availability measure-
Burt, W. H. 1943. Territoriality and home range concepts as applied to ments for evaluating resource preference. Ecology 61:65–71.
mammals. Journal of Mammalogy 24:346–352. Kapfer, J. M., J. R. Coggins, and R. Hay. 2008. Spatial ecology and habitat
Diemer, J. E., and D. W. Speake. 1983. The distribution of the eastern selection of bullsnakes (Pituophis catenifer sayi) at the northern periphery of
indigo snake, Drymarchon corais couperi, in Georgia. Journal of their geographic range. Copeia 2008:815–826.
Herpetology 17:256–264. Kramer, E. A., M. J. Conroy, M. J. Elliott, E. A. Anderson, W. R. Bumback,
Dodd, C. K., Jr. 1987. Status, conservation, and management. Pages 478– and J. Epstein. 2003. A GAP analysis of Georgia. Final report
513 in R. A. Seigel, J. T. Collins, and S. S. Novak, editors. Snakes: ecology and data. U.S. Department of Interior, U.S. Geological Survey.
and evolutionary biology. MacMillan, New York, New York, USA. Biological Resource Division, GAP Analysis Program, Athens, Georgia,
Dodd, C. K., Jr., and W. J. Barichivich. 2007. Movements of large snakes USA.
(Drymarchon, Masticophis) in north-central Florida. Florida Scientist 70: Lambeck, R. J. 1997. Focal species: a multi-species umbrella for nature
83–94. conservation. Conservation Biology 11:849–856.
Dunning, J. B., B. J. Danielson, and H. R. Pulliam. 1992. Ecological Landers, J. L., and D. W. Speake. 1980. Management needs of sandhill
processes that affect populations in complex landscapes. Oikos 65:169– reptiles in southern Georgia. Proceedings of the Annual Conference of
175. Southeast Association Fish and Wildlife Agencies 34:515–529.

110 The Journal of Wildlife Management  78(1)

Layne, J. N., and T. M. Steiner. 1984. Sexual dimorphism in occurrence of Shine, R. 2003. Reproductive strategies in snakes. Proceedings of the Royal
keeled dorsal scales in the eastern indigo snake (Drymarchon corais couperi). Society of London Series B Biological Sciences 270:995–1004.
Copeia 1984:776–778. Smith, C. R. 1987. Ecology of juvenile and gravid eastern indigo snakes in
Macartney, J. M., P. T. Gregory, and K. W. Larsen. 1988. A tabular study of north Florida. Thesis, Auburn University, Auburn, Alabama, USA.
data on movements and home ranges of snakes. Journal of Herpetology Southwood, T. E. 1966. Ecological methods: with particular reference to the
22:61–73. study of insect populations. Chapman & Hall, London, United Kingdom.
McCoy, E. D., and H. R. Mushinsky. 1999. Habitat fragmentation and the Speake, D. W. 1993. Indigo snake recovery plan revision. Final report to
abundances of vertebrates in the Florida scrub. Ecology 80:2526–2538. the U.S. Fish and Wildlife Service, Jacksonville, Florida, USA.
McNab, B. K. 1963. Bioenergetics and the determination of home range Speake, D. W., J. A. McGlincy, and T. R. Colvin. 1978. Ecology and
size. American Naturalist 97:133–140. management of the eastern indigo snake in Georgia: a progress report.
Mech, L. D. 1983. Handbook of animal radio-tracking. University of Pages 64–73 in R. R. Odum, and J. L. Landers, editors. Rare and
Minnesota Press, Minneapolis, USA. endangered wildlife symposium. Georgia Department of Natural
Mohr, C. O. 1947. Table of equivalent populations of North American small Resources, Game and Fish Division, Technical Bulletin WL 4, Social
mammals. American Midland Naturalist 37:223–249. Circle, USA.
Moler, P. E. 1985. Home range and seasonal activity of the eastern indigo Speake, D. W., J. A. McGlincy, and C. Smith. 1987. Captive breeding and
snake, Drymarchon corais couperi, in northern Florida. Final performance experimental reintroduction of the eastern indigo snake. Pages 84–88 in
report. Study E-1-06, III-A-5. Florida Game and Fresh Water Fish R. R. Odum, K. Riddleberger, and J. Ozier, editors. Third Southeast
Commission, Tallahassee, USA. Nongame and Endangered Wildlife Symposium. Georgia Department of
Moler, P. E. 1992. Eastern indigo snake. Pages 181–186 in P. E. Moler, Natural Resources, Game and Fish Division, Social Circle, USA.
editor. Rare and endangered biota of Florida: amphibians and reptiles. Steen, D. A., C. J. W. McClure, J. C. Brock, D. C. Rudolph, J. B. Pierce,
University Press of Florida, Gainesville, USA. J. R. Lee, W. J. Humphries, B. B. Gregory, W. B. Sutton, L. L. Smith,
Moyer, M. A., J. W. McCown, and M. K. Oli. 2007. Factors influencing D. L. Baxley, D. J. Stevenson, and C. Guyer. 2012. Landscape-level
home-range size of female Florida black bears. Journal of Mammalogy influences of terrestrial snake occupancy within the southeastern United
88:468–476. States. Ecological Applications 22:1084–1097.
Mushinsky, H. R., and E. D. McCoy. 1985. On the relationship between Stevenson, D. J., M. R. Bolt, D. J. Smith, K. M. Enge, N. L. Hyslop, T. M.
fire periodicity, plant structure and herpetofaunal communities in Florida. Norton, and K. J. Dyer. 2010. Prey records for the eastern indigo snake
American Zoologist 25:A9–A9. (Drymarchon couperi). Southeastern Naturalist 9:1–18.
Noss, R. F. 2000. High-risk ecosystems as foci for considering biodiversity Stevenson, D. J., K. J. Dyer, and B. A. Willis-Stevenson. 2003. Survey and
and ecological integrity in ecological risk assessments. Environmental monitoring of the eastern indigo snake in Georgia. Southeastern
Science and Policy 3:321–332. Naturalist 2:393–408.
Ott, P., and F. Hovey. 1997. Bycomp, Version 1.0. British Columbia Forest Stevenson, D. J., K. M. Enge, L. Carlile, K. J. Dyer, T. M. Norton, N. L.
Service, Victoria, Canada. < http://nhsbig.inhs.uiuc.edu/wes/habitat. Hyslop, and R. A. Kiltie. 2009. An eastern indigo snake (Drymarchon
html >. Accessed 5 Jan 2007. couperi) mark-recapture study in southeastern Georgia. Herpetological
Parker, W. S., and W. S. Brown. 1980. Comparative ecology of two colubrid Conservation and Biology 4:30–42.
snakes, Masticophis t. taeniatus and Pituophis melanoleucus deserticola in Tufto, J., R. Andersen, and J. Linnell. 1996. Habitat use and ecological
northern Utah. Milwaukee Public Museum Publications in Biology and correlates of home range size in a small cervid: the roe deer. Journal of
Geology, Milwaukee, Wisconsin, USA. Animal Ecology 65:715–724.
Pope, S. E., L. Fahrig, and N. G. Merriam. 2000. Landscape United States Fish and Wildlife Service [USFWS]. 1978. Endangered and
complementation and metapopulation effects on leopard frog populations. threatened wildlife and plants: listing of the eastern indigo snake as a
Ecology 81:2498–2508. threatened species. Federal Register 43:4026–4028.
Reiman, B. E., J. T. Peterson, and D. L. Myers. 2006. Have brook trout United States Fish and Wildlife Service [USFWS]. 2008. Eastern indigo
displaced bull trout in streams of central Idaho? An empirical analysis of snake (Drymarchon couperi) 5-year review: summary and evaluation.
distributions along elevation and thermal gradients. Canadian Journal of United States Fish and Wildlife Service, Southeast region, Mississippi
Fisheries and Aquatic Sciences 63:63–78. Ecological Services Field Office, Jackson, USA.
Reinert, H. K. 1984. Habitat variation within sympatric snake populations. Van Lear, D. H., W. D. Carroll, P. R. Kapeluck, and R. Johnson. 2005.
Ecology 65:1673–1682. History and restoration of the longleaf pine-grassland ecosystem:
Reinert, H. K. 1993. Habitat selection in snakes. Pages 201–240 in R. A. implications for species at risk. Forest Ecology and Management
Seigel, and J. T. Collins, editors. Snakes: ecology and behavior. McGraw- 211:150–165.
Hill, Inc., New York, New York, USA. Waldron, J. L., S. M. Welch, and S. H. Bennett. 2008. Vegetation structure
Reinert, H. K., and R. T. Zappalorti. 1988. Timber rattlesnakes (Crotalus and the habitat specificity of a declining North American reptile: a
horridus) of the pine barrens: their movement patterns and habitat remnant of former landscapes. Biological Conservation 141:2477–2482.
preference. Copeia 1988:964–978. Wharton, C. H. 1978. The natural environment of Georgia. Georgia
Roberge, J. M., and P. Angelstam. 2004. Usefulness of the umbrella species Department of Natural Resources, Atlanta, USA.
concept as a conservation tool. Conservation Biology 18:76–85. Whitaker, P. B., and R. Shine. 2003. A radiotelemetric study of movements
Samuel, M. D., D. J. Pierce, and E. O. Garton. 1985. Identifying areas of and shelter-site selection by free-ranging brownsnakes (Pseudonaja textilis,
concentrated use within the home range. Journal of Animal Ecology Elapidae). Herpetological Monographs 17:130–144.
54:711–719. Worton, B. J. 1989. Kernel methods for estimating the utilization
SAS Institute, Inc. 2005. Version 9.1 user manual. SAS Institute, Cary, distribution in home range studies. Ecology 70:164–168.
North Carolina, USA.
Seaman, D. E., and R. A. Powell. 1996. An evaluation of the accuracy of
kernel density estimators for home range. Ecology 77:2075–2085. Associate Editor: Carola Haas.

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