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2015 SOUTHEASTERN NATURALIST 14(2):342–350
Spatial Ecology and Habitat Use of the Coachwhip in a
Longleaf Pine Forest
Jennifer M. Howze1,* and Lora L. Smith1
Abstract - We examined spatial ecology and habitat use of Coluber flagellum (Coachwhip)
in a 12,000-ha Pinus palustris (Longleaf Pine) reserve in southwestern Georgia from 2007
through 2008. We radio-tracked 7 Coachwhips (5 males and 2 females) for 291 to 325 days.
The average 100% minimum convex polygon (MCP) home-range for all snakes was 102.9
± 28 ha. Daily movement during the active season (April–November) varied from 28.6 to
73.6 m for males (n = 5) and from 27.5 to 95.6 m for females (n = 2). Snakes were usually
associated with open-canopied pine forests and found less often in aquatic and agricultural
habitats. Our results are consistent with evidence from previous studies in that Coachwhips
used sites with open-forest structure and large expanses of habitat.
Introduction
Knowledge of how an animal uses space is vital to understanding its ecology
(Gregory et al. 2001). Many factors, including habitat structure and reproduction
(Gibbons and Semlitsch 2001, Gregory et al. 2001), prey availability (King and
Duvall 1990, Shine et al. 2003), competition (Moore 1978), predator density (Shine
and Lambeck 1985), and environmental conditions (Lillywhite 2001, Webb and
Shine 1998), influence spatial ecology and movement of snakes and play a role
in determining activity levels. Foraging ecology may also be an important indicator
in predicting spatial-use and movement patterns for wide-ranging species.
The most-common foraging modes exhibited by squamates fall within 2 general
categories: sit-and-wait foragers that ambush active prey, and active foragers, like
Coluber flagellum Shaw (Coachwhip), that hunt active and sedentary prey as they
move through the landscape (Cooper and Whiting 2000, Huey and Pianka 1981,
Mushinsky 2001, Schoener 1971, Secor 1995). The active-foraging strategy balances
higher predation risk and greater energy expenditure with increased energy
acquisition through high food intake (Secor 1995, Secor and Nagy 1994) and can
result in species like Coachwhips potentially traveling great distances (>1 km per
day) and using large home-ranges (McCartney et al. 1988, Secor 1995).
The Coachwhip has an expansive geographic range that extends from North
Carolina to southern Florida and west to Texas, Oklahoma, and southeastern Kansas
(Conant and Collins 1998). In the Southeast, Coachwhips are strongly tied to
the southeastern coastal plain, which is characterized by xeric upland habitats (Tuberville
and Gibbons 2008) and was once dominated by the currently endangered
Pinus palustris Mill (Longleaf Pine) ecosystem (Edwards et al. 2013). Although
1Joseph W. Jones Ecological Research Center, 3988 Jones Center Drive, Newton, GA
39870. *Corresponding author - jhowze@jonesctr.org.
Manuscript Editor: Natalie Hyslop
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Coachwhips are not endemic to the Longleaf Pine ecosystem (Halstead et al. 2009,
Johnson et al. 2007, Mitrovich et al. 2009), remnants of this once wide-ranging
ecosystem likely play a significant role in the species’ persistence. Few studies
(Baxley and Qualls 2009, Dodd and Barichivich 2007) have quantified spatial ecology
and habitat use of Coachwhips within the Longleaf Pine ecosystem. Therefore,
we present radio-telemetry data to address informational gaps on how Coachwhips
use components of this important habitat in southwestern Georgia.
Methods
This study was conducted at Ichauway, the research site of the Joseph W. Jones
Ecological Research Center in Newton, GA. The 12,000-ha site consisted mainly
of second-growth Longleaf Pine savanna managed on a 2-year prescribed-fire rotation.
Patches of closed-canopied Quercus spp. (oak) forests occurred primarily
in isolated depressions, around seasonally inundated wetlands, and along 45 km
of Ichawaynochaway Creek and the Flint River. Wildlife-food plots (comprised of
Sorghum bicolor [Milo], Triticum spp. [wheat], and Zea mays [Corn]) and Longleaf
Pine plantations were scattered throughout the property. The property was surrounded
by center-pivot-irrigated agricultural lands.
We captured snakes using 16 box-trap arrays (Burgdorf et al. 2005) located in
Longleaf Pine savanna habitat with native groundcover species, including Aristida
stricta Michx. (Wiregrass), Andropogon spp. (broomsedge), and Pteridium aquilinum
(L.) Kuhn (Bracken Fern). We checked the traps 3 times per week from March
through November in 2007 and 2008.
For each snake captured, we collected snout-to-vent length (SVL), tail length,
and body mass measurements and identified sex through cloacal probing. We
marked captured snakes using passive integrated transponder (PIT) tags injected
subcutaneously between the dorsal and ventral scales on the lower third of the body
(Gibbons and Andrews 2004). We selected 5 adult male and 5 adult female Coachwhips
based on size (>74 cm SVL) and sex (we attempted to maintain an equal sex
ratio) and surgically implanted them with 9-g radio transmitters with an 18-month
battery life (model SI-2; Holohil Systems Ltd., Carp, ON, Canada) using methods
described in Reinert and Cundall (1982). We radio-tracked Coachwhips (SVL range
= 134.7–160.4 cm) for 11 months, from June 2007 through April 2008. We located
snakes 1–2 times per week using triangulation techniques (≥2 bearings collected)
during the active season (April–November) because snakes were often moving as
we tracked them. We honed in on snake locations using radio telemetry during the
inactive season (December–March) when snakes were overwintering. We minimized
triangulation error by discarding bearings under the following conditions if:
(1) signal strength was weak, (2) the angle between locations was <45° or >135°,
or (3) the sequential bearing observations were greater than 15 min apart (White
and Garrott 1990, Withey et al. 2001). We recorded all snake locations (UTM coordinates)
on a handheld PDA accurate to within 3 m (Garmin IQue 3600; Garmin
International, Inc., Olathe, Kansas) and calculated triangulated locations using Program
Locate software (Nams 2006; Pacer Computing, Tatamagouche, NS, Canada).
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We employed ArcGIS 9.3.1 (Environmental Systems Research Institute, Redlands,
CA) to create a spatial layer that included snake locations.
We used Hawth’s Tools extension (Beyer 2004) in ArcGIS 9.3.1 to construct
100% minimum convex polygon (MCP; Mohr 1947) home-ranges and Home-Range
Tools extension in ArcGIS 9.3.1 to calculate 95% and 50% (core) MCPs to facilitate
comparison with other published studies. We used linear regression to identify if
length of tracking period or number of tracking events were related to home-range
size (100% MCPs). We used Hawth’s Tools extension to calculate the distance
between consecutive points for each snake and standardized distance estimates by
calculating daily movements (straight-line distance between consecutive points divided
by the number of days between tracking events).
We created a spatial-data layer using ArcGIS 9.3.1 that included unique
snake locations plotted within an existing land-cover (habitat) data layer that
was digitized using 1:12,000-scale color infrared aerial photography and from
ground truthing observations. The land-cover layer included 4 habitat classes:
(1) agriculture/scrub habitat (wildlife food plots and old fields); (2) hardwood
forest (Quercus falcata Michx. [Southern Red Oak], Q. virginiana Mill. [Live
Oak], Q. laurifolia Michx. [Laurel Oak], and Q. nigra L. [Water Oak]); (3) pine
forest (natural Longleaf Pine savanna), Longleaf Pine plantation (sawtimber- and
pole-size classes), and mixed natural pine forests (50–80% Longleaf Pine savanna
mixed with oaks); and (4) aquatic habitat (isolated wetlands and Ichawaynochaway
Creek). We defined the study area as a 100% MCP for all snake locations
with a 500-m buffer, which we considered to include habitat available to snakes
based on the daily distance Coachwhips traveled during this study (averages all
<100 m).
We used compositional analysis (Aebischer et al. 1993) to test for second-order
(landscape level) and third-order (home-range level) habitat use (Johnson 1980).
To test for habitat use at the home-range scale, we used multivariate analyses of
variance (MANOVA) to compare (1) habitat use at snake locations to habitat available
within home range (100% MCP), (2) habitat use at snake locations to habitat
available at the core area (50% MCP), and (3) available habitat within the core area
(50% MCP) to available habitat within the home range (100% MCP). To test for
habitat use at the landscape scale, we compared available habitat within the home
range (100% MCP) to available habitat within the study area.
Results
We radio tracked 7 Coachwhips (5 males and 2 females) from 291 to 325 days.
Two of the 10 snakes implanted with radio-transmitters died (1 female unknown
mortality, 1 female egg-bound), and a third snake was lost within the first 2 months
of the study; we excluded these 3 individuals from analyses. We observed an
average of 27 unique locations per snake (range = 18–31) and an average of 40
tracking events (includes locations where snakes remained in the same location
during the inactive season) per snake (range = 27–47). During our sampling period
from June 2007 through April 2008, snakes exhibited the greatest daily movement
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during June–September and the following April (Fig. 1). We observed decreased
daily movement in October and November, and did not observe movement for most
snakes (6 of 7) during the coldest months (December–February). Average daily
movement of snakes during the active season (April–November, Fig. 1.) varied
from 28.6 to 73.6 ± 27.8 m for males (n = 5) and 27.5–95.6 ± 22.1 m for females
(n = 2). Six of 7 snakes had daily movements that exceeded 100 m (15.6% of active
season locations), and we recorded a maximum daily movement of 224 m made by
a female.
Home-range size estimates were not significantly correlated with tracking
period (R² = 0.10, P = 0.51) or number of tracking events (R² = 0.21, P = 0.30).
Average 100% MCP home range for all snakes was 102.9 ± 28 ha (Table 1.). Male
snakes had an average 100% MCP that was more than twice that of female snakes
Figure 1. Average daily movement of male (n = 5) and female (n = 2) Coluber flagellum
(Coachwhip) radio-tracked from June 2007 through April 2008 in Baker County, GA.
Table 1. Average home-range estimates (ha) using minimum convex polygons (MCP) for 7 telemetered
Coluber flagellum (Coachwhip) radio-tracked from June 2007 to April 2008 in Baker County,
GA.
MCP 100% MCP 95% MCP 50%
All snakes
Average (SD) 102.9 (28.0) 84.9 (32.0) 12.7 (9.1)
Range 59.1–132.6 48.1–132.6 3.9–28.9
Male (n = 5)
Average (SD) 117.9 (13) 98.2 (27.5) 12.8 (10.0)
Range 113.0–132.6 58.0–132.6 3.9–28.9
Female (n = 2)
Average (SD) 65.2 (9) 51.7 (5.0) 12.3 (10.1)
Range 59.1–71.3 48.1–55.2 5.2–19.4
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(males: 117.9 ± 12.8 ha, females: 65.2 ± 8.6 ha). Average 50% MCP core areas were
similar in size for the following: all snakes: 12.7 ± 9.1 ha, males: 12.8 ± 10.0 ha,
and females: 12.3 ± 10.1 ha.
Coachwhips used habitats relative to their availability at the landscape scale
(F = 2.75, P = 0.21), home-range scale (F = 0.06, P = 0.98), and the core-area
scale (F = 4.42, P = 0.13); however, composition of habitat within the core home
range differed significantly from that of the 100% MCP home range (F = 16.3, P =
0.02). Specifically, the proportion of pine forests in core areas was greater than
expected based on availability within the 100% MCP, whereas the proportion of
aquatic and agricultural habitats in core areas were less than expected (Fig. 2).
Discussion
Studies have suggested that large-bodied terrestrial snakes require extensive
habitat to maintain their populations (Dodd and Barichivich 2007, Hyslop et al.
2014, Mitrovich 2006). Previous research at our study site found that average homeranges
for large-bodied sit-and-wait foragers, including Pituophis melanoleucus
Daudin (Pinesnake; 100% MCP = 59.2 ha; Miller et al. 2012), Lampropeltis getula
L. (Eastern Kingsnake; 100% MCP = 49.5 ha; Linehan et al. 2010), and Crotalus
adamanteus Palisot de Beauvois (Eastern Diamondback Rattlesnake; 100% MCP
= 24.6 ha; Hoss et al. 2010), were smaller in comparison to that of the Coachwhip.
Figure 2. Average proportional core-habitat use by Coluber flagellum (Coachwhip) relative
to availability (home range) in Baker County, GA.
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Coluber constrictor L. (North American Racer), a congener of the Coachwhip that
exhibits a similar active-foraging mode but a smaller body size, reportedly has
smaller average MCP home ranges (11.45 ha and 12.2 ha, respectively; Klug et
al. 2011, Plummer and Congdon 1994) than that of the Coachwhip. Another active
forager, Drymarchon couperi Holbrook (Eastern Indigo Snake), one of the largest
native snakes in the Southeast, used larger average home-ranges (100% MCP > 340
ha) than did Coachwhips in similar habitats (Hyslop et al. 2014). Therefore, a combination
of an active-foraging strategy and a large body size may be important in
explaining the larger spatial requirements for snakes like Coachwhips and Eastern
Indigo Snakes.
We observed long-distance daily movements exceeding 100 m for 6 of 7 snakes
during the active season. We assume that we missed some additional long-distance
movements in our study because we tracked snakes once per week and we were
unable to sample during May, when Coachwhips were most active at our study
site (J.M. Howze, unpubl.data). Additionally, we calculated our distance estimates
as straight-line measurements between locations, which likely underestimated the
actual length of paths traveled by snakes. Nonetheless, our findings, along with
research on Coachwhips in Texas (100% MCP = 70.4 ha; Johnson et al. 2007),
California (100% MCP = 136.4 ha; Mitrovich et al. 2009), and Florida (100% MCP
= 183 ha [males], 102 ha [females]; Halstead et al. 2009) support the body of evidence
describing the large spatial requirements necessary for this species across
its range. Secor (1995) found that these large home-ranges reflected frequent longdistance
movements by Coachwhips.
Previous studies have reported that Coachwhips were found in a variety of
open-canopy, xeric, southeastern forest types including Longleaf Pine, scrub,
oak savanna, sandhills, and pine flatwoods (Dodd and Barichivich 2007, Halstead
et al. 2009, Johnson et al. 2007, Tuberville and Gibbons 2008). Our data
suggested that Coachwhips used pine forest more often in their core areas and
were less likely to use aquatic and agricultural habitats, suggesting that habitat
structure might be an important variable in explaining habitat selection in
Coachwhips. Further evidence provided by Baxley and Qualls (2009) described
a positive correlation between Coachwhips and xeric open-canopy areas within
Longleaf Pine habitats.
Coachwhip foraging strategy may explain a propensity for open-forest structure.
They are visual predators, and areas with sparse vegetation may be helpful for hunting
(Ernst and Ernst 2003) lizards (their primary prey) and small mammals (Halstead
et al. 2008, Hamilton and Pollack 1956, Secor 1995). Furthermore, Coachwhips use
structural features of open-canopy habitats like rotting pine stumps, root holes, and
animal burrows to forage for prey, escape predators, and regulate body temperature
during thermal extremes (Dodd and Barichivich 2007, Ernst and Ernst 2003, Gentry
and Smith 1968, Secor 1995, Secor and Nagy 1994, Tuberville and Gibbons 2008).
Forest-management practices such as prescribed fire and thinning, which maintain an
open-canopy structure, and the protection of contiguous habitat may help to provide
appropriate habitat for Coachwhips in the Southeast.
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Acknowledgments
Funding for the project was provided by the Florida Fish and Wildlife Conservation
Commission’s Wildlife Legacy Initiative program and the US Fish and Wildlife Service’s
State Wildlife Grants program (Grant # SWG 05-020, Agreement #060010). We thank Kelly
McKean, Aubrey Heupel, Stephen Jones, Phil Shirk, Chris Thawley, and Billy Thein for
field assistance, and Terry Norton, DVM, for surgically implanting radio-transmitters into
our study animals.
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