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Home Range, Survival, and Activity Patterns of the
Southeastern Pocket Gopher: Implications for
Translocation
Ashley E. Warren, L. Mike Conner, Steven B. Castleberry,* Daniel Markewitz
A.E. Warren, S.B. Castleberry, D. Markewitz
Warnell School of Forestry and Natural Resources, University of Georgia, Athens, Georgia 30602
Present address of A.E. Warren: Florida Fish and Wildlife Conservation Commission, Panama City, Florida 32409
L.M. Conner
Joseph W. Jones Ecological Research Center at Ichauway, Newton, Georgia 39870
Abstract
The southeastern pocket gopher Geomys pinetis is absent from a large portion of its historical range. Translocation may
represent a viable management technique to reestablish populations into suitable habitat. However, several aspects of
the species’ ecology are poorly understood, making development of an effective translocation approach challenging.
Therefore, we used radiotelemetry to examine home range, survival, dispersal, and daily activity patterns of the
southeastern pocket gopher in southwestern Georgia. We measured soil and vegetation characteristics within
individual home ranges and examined relationships between home range size, habitat variables, and body mass. Mean
home range size of 17 radio-tagged pocket gophers was 921.9 m
2
(range ¼43.4–2246.8 m
2
). Home range size was
positively related to body mass, percent silt at a depth of 25 cm, and soil carbon content at 75 cm and was negatively
related to percent sand at 25 cm, percent clay at 50 cm, and ground cover of grasses other than wiregrass Aristida
beyrichiana. Survival rate was 0.78 (range ¼0.50–1.00) over the 51-wk study, and we documented predation, likely by
avian predators, on two individuals. Three individuals dispersed, with a maximum dispersal distance of 319.1 m (range
¼143.2–319.1 m), the farthest known southeastern pocket gopher dispersal. Pocket gophers exhibited greater activity
from 0000 to 0400 hours and from 1600 to 2000 hours, contrasting previous research that southeastern pocket
gophers were equally active throughout the diel period. Our home range size estimates for southeastern pocket
gophers are the first determined using radiotelemetry and are considerably smaller than previous estimates. Although
we documented dispersal distances more than 300 m, the fragmented nature of current and restored habitats likely
will prevent large-scale natural colonization. Our results provide information important for maximizing success in
future southeastern pocket gopher translocation efforts.
Keywords: Geomys pinetis; southeastern pocket gopher; home range; survival; activity patterns; translocation
Received: March 10, 2017; Accepted: September 20, 2017; Published Online Early: September 2017; Published:
December 2017
Citation: Warren AE, Conner LM, Castleberry SB, Markewitz D. 2017. Home range, survival, and activity patterns of the
southeastern pocket gopher: Implications for translocation. Journal of Fish and Wildlife Management 8(2):544-557;
e1944-687X. doi:10.3996/032017-JFWM-023
Copyright: All material appearing in the Journal of Fish and Wildlife Management is in the public domain and may be
reproduced or copied without permission unless specifically noted with the copyright symbol &. Citation of the
source, as given above, is requested.
The findings and conclusions in this article are those of the author(s) and do not necessarily represent the views of the
U.S. Fish and Wildlife Service.
* Corresponding author: scastle@warnell.uga.edu
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 544
Introduction
The southeastern pocket gopher Geomys pinetis was
historically associated with the longleaf pine Pinus
palustris community characteristic of the Coastal Plain
physiographic province in southeastern Alabama, south-
ern Georgia, and northern and central Florida (Golley
1962; Pembleton and Williams 1978; Wilkins 1987).
Longleaf pine forests have been highly impacted by
conversion and fragmentation, resulting in habitat loss
and alteration for many associated species, including the
southeastern pocket gopher. Although southeastern
pocket gophers can be abundant in suitable habitat,
the species is absent from a large portion of its historical
range (Georgia Department of Natural Resources 2005).
As a result, Alabama, Georgia, and Florida state wildlife
agencies have listed the southeastern pocket gopher as
a high priority species in their State Wildlife Action Plans
(Alabama Department of Conservation and Natural
Resources 2005; Georgia Department of Natural Resourc-
es 2005; Florida Fish and Wildlife Conservation Commis-
sion 2012). Species in this category show combinations
of rarity, limited distribution, decreasing size or viability
of populations, and biological vulnerability.
Developing a conservation strategy for any species
requires a basic understanding of its ecology. However,
several aspects of southeastern pocket gopher ecology
are poorly understood, primarily because its fossorial
lifestyle makes observational studies difficult. The only
existing home range estimates are based on a single
study that determined maximum dimensions of mound
patterns (Hickman and Brown 1973a) that likely included
considerable unused area and subsequently overesti-
mated home range sizes. Although southeastern pocket
gopher home range data are lacking, information is
available for other Geomys species. Cameron et al. (1988)
concluded that the area covered by the burrow systems
of Attwater’s pocket gopher Geomys attwateri was highly
variable between individuals, and uncorrelated differ-
ences with sex, age, or body size. Conversely, home
range size of the Ozark pocket gopher Geomys bursarius
ozarkensis was directly proportional to body size in
juvenile females, inversely proportional to body size in
adult females, and uncorrelated with body size in males
(Connior and Risch 2010). Although presence of south-
eastern pocket gophers is influenced by soil (Warren et
al., in press) and vegetation (Ross 1976) characteristics,
how those habitat features influence home range size is
unknown. Given the variability within and among
studies, generalizing results from other Geomys species
to the southeastern pocket gopher could lead to
erroneous conclusions.
Southeastern pocket gophers likely have a high
survival rate due to the protection burrows provide
from predators. Brown (1971) suggested that longevity
of southeastern pocket gophers in Florida was more than
2 y. The only published study that used radiotelemetry to
investigate survival rates in pocket gophers occurred on
the Ozark pocket gopher in north central Arkansas
(Connior and Risch 2010). They reported that 33 of 35
pocket gophers survived over 144 d during the
nonbreeding season and 26 of 35 survived over 116 d
during the breeding season. Literature regarding cause-
specific mortality in pocket gophers is equally sparse.
Connior and Risch (2010) attributed 7 of 11 Ozark pocket
gopher mortalities to predation, but the predator could
only be identified in a single case when a tagged
individual was predated by a prairie kingsnake Lamro-
peltis calligaster calligaster. The Florida pine snake
Pituophis melanoleucus mugitus is likely the most
common predator on southeastern pocket gophers
due to its presence in the same habitats and its ability
to exploit fossorial prey (Miller et al. 2012).
Because suitable habitat currently exists in fragmented
patches (Georgia Department of Natural Resources
2005), information on dispersal periodicity, timing, and
distance is needed to determine whether dispersal
behavior would facilitate establishment of southeastern
pocket gopher populations in patches of suitable
habitat. If not, a successful conservation strategy may
include reestablishment of populations through translo-
cation. Although fragmentation may be limiting natural
dispersal, information on dispersal behavior is limited to
anecdotal reports in which two southeastern pocket
gophers dispersed 184 and 244 m, respectively (Hickman
and Brown 1973a). Dispersal has been observed in
Botta’s pocket gopher Thomomys bottae, with most
dispersal activity occurring during the spring and
summer before reproductive age (Daly and Patton
1990). Whether the same pattern occurs for southeastern
pocket gophers is unknown.
The only published research directly addressing daily
activity patterns in southeastern pocket gophers was
conducted in captivity as part of a thermoregulation
study. Ross (1980) concluded that southeastern pocket
gophers alternate periods of activity throughout the day
and night in roughly 40-min cycles. Although the
method for measuring activity in the study was precise,
activity may have been influenced by the inability to
exhibit natural foraging behavior. In contrast, anecdotal
field observations suggest that most mounding activity
occurs at dusk and dawn (Hickman and Brown 1973a).
Mounding activity may be useful as a proxy for activity;
however, other daily activities may occur completely
below ground and would not be detected through
mounding activities alone.
Further loss of the southeastern pocket gopher from
its historic range may have profound negative effects on
upland ecosystems of the southeastern Coastal Plain,
because they play vital roles in the communities they
inhabit. Their mounds are the most common source of
faunal soil disturbance within longleaf pine communities
(Simkin and Michener 2005). In the Sandhills ecosystem
of the southeastern Coastal Plain, several species of
amphibians and reptiles use southeastern pocket gopher
mounds as shelter (Funderburg and Lee 1968), including
the gopher frog Lithobates capito (Blihovde 2006) and
mole skink Plestiodon egregius (Mount 1963). The
mounds and tunnels also serve as habitat for several
arthropods, many of which are believed to be obligate
commensals (Pembleton and Williams 1978; Skelley and
Kovarik 2001). Given their importance in maintaining the
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 545
integrity of these systems, an effective conservation
strategy is needed, yet the current information regarding
several aspects of their natural history is insufficient.
Therefore, we used radiotelemetry to investigate home
range, survival, cause-specific mortality, dispersal, and
daily activity patterns. Information resulting from this
study will be integral in forming an effective conserva-
tion strategy to restore the species into suitable habitat
within its historic range.
Methods
Study site
We conducted our study from September 2012 to
September 2013 at the Joseph W. Jones Ecological
Research Center at Ichauway in Baker County, Georgia
(Figure 1). Ichauway covers 117 km
2
of predominately
longleaf pine forest surrounded primarily by agriculture.
Other cover types include slash pine Pinus elliottii and
loblolly pine Pinus taeda forests, mixed pine–hardwoods,
riparian hardwood forests, live oak Quercus virginianus
depressions, isolated wetlands, creek swamps, agricul-
tural fields, and areas impacted by human development.
Wiregrass Aristida beyrichiana is the dominate understory
species covering approximately one-third of the proper-
ty. Habitat structure and composition are maintained
through prescribed fire. Stands are burned at least every
other year, primarily during March and April (Atkinson et
al. 1996). Ichauway is located in the Dougherty Plain
physiographic district that is characterized by marine and
fluvial deposited parent material that now comprise
Entisols and Ultisols over highly fractured Ocala lime-
stone, and a flat to rolling karst topography (Beck and
Arden 1984; Hayes et al. 1983; Couch et al. 1996).
Animal capture and transmitter implantation
We selected locations for pocket gopher trapping
through opportunistic sightings of mounds. We main-
tained more than 250 m between radio-tagged individ-
uals, which represented the furthest documented
dispersal of southeastern pocket gophers prior to our
study. We captured pocket gophers by placing live traps
described by Hart (1973) and Connior and Risch (2009a)
in excavated tunnels. We checked traps every 3 h. We
placed captured individuals in ventilated 45.4-L plastic
containers partially filled with moist soil from the site and
transported them to the lab for transmitter implantation.
We surgically implanted 3-g VHS radio transmitters
(SOPI-2070, Wildlife Materials Inc., Murphysboro, IL),
representing a mean of 1.73% (standard deviation [SD]
¼0.48, range ¼0.89–2.38) of body mass, either
subcutaneously between the scapulae or within the
peritoneal cavity of subadult and adult pocket gophers
(body mass .100 g; Wing 1960). We inserted a passive
integrated transponder tag subcutaneously away from
the surgery site. Transmitter and PIT tag implantation
occurred under continuously inhaled sevoflurane. We
held gophers 3 d postsurgery to monitor recovery and
manage inflammation and pain with intramuscular
injections of meloxicam and butorphanol. We provided
gophers with sweet potatoes from the time of capture
until returned to their original burrows. We did not
replace failed or lost transmitters. Animal capture and
transmitter implantation followed guidelines of the
University of Georgia Animal Welfare Assurance A3437-
01 and were approved by the Institutional Animal Care
and Use Committee of the University of Georgia (Animal
Use Proposals 04-002-Y1-A0, A2012 04-002-A1, A2012
04-002-R1).
Radio tracking
We determined the location of radio-tagged pocket
gophers during tracking periods conducted every other
day by using a telemetry receiver (Communication
Specialists, Inc., R-1000) and three-element Yagi antenna.
We began each tracking period 2 h later than the
previous to account for activity around the diel period.
We began tracking outside the known area of gopher
activity and homed in on the point of greatest signal
strength with the antenna pointed downward, being
careful to limit ground vibration while walking to avoid
influencing gopher movements. We recorded one
location for each individual during each tracking period
by using a NomadtGlobal Positioning System (GPS;
Trimble Navigation, Ltd., Sunnyvale, CA) equipped with a
GPS antenna (Crescent A100, Hemisphere GPS, Inc.,
Mountain View, CA) that provided a horizontal accuracy
of less than 0.6 m with 95% confidence. In cases of
dispersal, we continued tracking the dispersed individual
at its new location. We defined dispersal as complete
abandonment of one area of concentrated activity to a
new area where the individual had not previously been
recorded.
We tracked individual pocket gophers until the
transmitter failed, mortality occurred, or the individual
could not be relocated. We attempted to relocate lost
individuals by first covering as much of the immediately
surrounding area by foot as could be traversed by one
person in 1–2 h. We then spent another 2–3 h using
property roads to search the area by vehicle, spanning
outwards from the last known location of the individual.
We confirmed transmitter failure either by a marked
decrease in signal strength before loss of signal, or by
recapturing the animal. In cases of mortality, we
determined cause of death by investigating conditions
at the mortality site, and by performing a necropsy.
When we determined predation as the cause of death,
we identified the suspected predator to the lowest taxon
possible based on carcass condition, tooth marks,
surrounding prints, and scat identification.
We conducted focal telemetry (extended tracking
periods focusing on fine-scale movements of an individ-
ual study animal) for nine pocket gophers from May
through August 2013 to investigate daily activity
patterns. Focal telemetry was conducted by tracking
each individual for 8 h/d on three separate days, all
conducted within a 2-wk period. Each day’s tracking
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 546
session covered a different 8-h portion of the 24-h diel
period. During each 8-h tracking session, we located the
gopher every 20 min. We recorded locations using a GPS
as described above. We quantified daily activity by
recording the distance traveled in each 20-min interval
between recordings and the distance from the nest at
each location. We determined distance traveled by
measuring between each sequential location by using
a meter tape. We used the Near tool in ArcMAP 9.3.1 to
measure distance between each recorded location and
the location of the nest. We assumed pocket gopher
nests were located where telemetry point density was
highest. To standardize for individual activity differences,
we converted distances to proportions by dividing each
measured distance by the longest distance recorded for
that individual during the 24 h of focal telemetry.
Vegetation and soil sampling
To investigate associations between home range size
and habitat features, we quantified vegetation and soil
characteristics within the home range of each radio-
tracked individual (Table 1). At each site, we randomly
selected five 1-m
2
subplots within 18 m of the center of
the home range. We quantified vegetation structure by
visually estimating percent ground cover of pine litter,
hardwood leaf litter, woody vegetation, forbs and vines,
Figure 1. Historic range of the southeastern pocket gopher Geomys pinetis and location of the Joseph W. Jones Ecological Research
Center at Ichauway in Baker County, Georgia.
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 547
wiregrass, and other grass species in each quadrant of
the subplot and averaging the quadrants.
We used a 7-cm-diameter bucket auger to collect soil
samples at depths of 0–10, 15–25, 40–50, 65–75, and 90–
100 cm at the center of each home range. We used the
qualitative field texture method to estimate soil texture
at each depth (Thien 1979) and used the soil textures at
each depth to create a texture profile for each site.
Selected representatives of each unique texture profile
were quantified at a commercial testing laboratory
(Waters Agricultural Testing Lab, Camilla, GA) by using
the hydrometer method for determining percent sand,
silt, and clay (Gee and Bauder 1986). We assigned the
quantified results of each representative profile to the
remaining samples from the sites that shared the same
profile. Percent nitrogen and carbon of each soil sample
were determined using a Flash 2000 carbon nitrogen
analyzer (CE Elantech, Lakewood, NJ) at the University of
Georgia Forest Soil Laboratory (Athens, GA). We deter-
mined pH for each sample by combining 5 g of soil with
10 mL of deionized water and immersing an electronic
pH probe in the solution (McLean 1982).
Data analysis
Due to the difficulty in determining sex of pocket
gophers based on external morphology (Baker et al.
2003), we pooled all individuals for analysis. We created
an area-observation curve using the bootstrap function
in R 2.3.1 (R Foundation for Statistical Computing) to
determine the minimum number of locations required to
estimate an accurate home range (Odum and Kuenzler
1955). We used a less than 5% increase to indicate the
asymptote (Laundr´
e and Keller 1984; Springer 2003) to
prevent excluding an excessive number of individuals
and overly decreasing sample size. We created minimum
convex polygons for each pocket gopher that met the
Table 1. Ground cover and soil texture variables (abbreviations in parentheses) within 18 m of the center of radio-tagged
southeastern pocket gopher Geomys pinetis home ranges measured as part of a study examining home range, survival, and activity
patterns in Baker County, Georgia, 2012–2013.
Variable Description
Vegetation ground cover
Pine straw (PS) Percent area of five 1-m
2
plots covered by pine straw
Leaf litter (LL) Percent area of five 1-m
2
plots covered by hardwood leaf litter
Woody vegetation (WO) Percent area of five 1-m
2
plots covered by woody vegetation
Forbs/vines (FV) Percent area of five 1-m
2
plots covered by forbs and vines
Wiregrass (WG) Percent area of five 1-m
2
plots covered by wiregrass
Other grass (OG) Percent area of five 1-m
2
plots covered by other grasses
Soil texture
Sand at 10 cm (SA) Percent sand of soil samples collected at 10 cm
Sand at 25 cm (SB) Percent sand of soil samples collected at 25 cm
Sand at 50 cm (SC) Percent sand of soil samples collected at 50 cm
Sand at 75 cm (SD) Percent sand of soil samples collected at 75 cm
Sand at 100 cm (SE) Percent sand of soil samples collected at 100 cm
Silt at 10 cm (TA) Percent silt of soil samples collected at 10 cm
Silt at 25 cm (TB) Percent silt of soil samples collected at 25 cm
Silt at 50 cm (TC) Percent silt of soil samples collected at 50 cm
Silt at 75 cm (TD) Percent silt of soil samples collected at 75 cm
Silt at 100 cm (TE) Percent silt of soil samples collected at 100 cm
Clay at 10 cm (CA) Percent clay of soil samples collected at 10 cm
Clay at 25 cm (CB) Percent clay of soil samples collected at 25 cm
Clay at 50 cm (CC) Percent clay of soil samples collected at 50 cm
Clay at 75 cm (CD) Percent clay of soil samples collected at 75 cm
Clay at 100 cm (CE) Percent clay of soil samples collected at 100 cm
Soil chemistry
Nitrogen at 10 cm (NA) Percent nitrogen of soil samples collected at 10 cm
Nitrogen at 25 cm (NB) Percent nitrogen of soil samples collected at 25 cm
Nitrogen at 50 cm (NC) Percent nitrogen of soil samples collected at 50 cm
Nitrogen at 75 cm (ND) Percent nitrogen of soil samples collected at 75 cm
Nitrogen at 100 cm (NE) Percent nitrogen of soil samples collected at 100 cm
Carbon at 10 cm (RA) Percent carbon of soil samples collected at 10 cm
Carbon at 25 cm (RB) Percent carbon of soil samples collected at 25 cm
Carbon at 50 cm (RC) Percent carbon of soil samples collected at 50 cm
Carbon at 75 cm (RD) Percent carbon of soil samples collected at 75 cm
Carbon at 100 cm (RE) Percent carbon of soil samples collected at 100 cm
pH at 10 cm (PA) pH of soil samples collected at 10 cm
pH at 25 cm (PB) pH of soil samples collected at 25 cm
pH at 50 cm (PC) pH of soil samples collected at 50 cm
pH at 75 cm (PD) pH of soil samples collected at 75 cm
pH at 100 cm (PE) pH of soil samples collected at 100 cm
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 548
required number of locations by using the Hawth’s tools
extension in ArcMAP 9.3.1. For individuals that dispersed,
we created the minimum convex polygon for the area
with the greatest number of locations.
We conducted correlation analyses using the CORR
procedure in SAS 9.3 (SAS Institute, Inc., Cary, NC) to
determine relationships of home range size with body
mass and habitat variables. We used Pearson’s product
moment correlation for normally distributed variables
and Spearman’s rank correlation for non-normal vari-
ables. We created a survivorship curve by using the
Kaplan–Meier Staggered Entry method (Pollock et al.
1989) based on weekly counts of individuals added to
the study, lost to mortality, or censored (i.e., removed
because of transmitter failure or because fate of the
individual was unknown). We described instances of
mortality and dispersal anecdotally due to a low number
of occurrences.
We separated daily activity data into six 4-h segments
and treated the proportional distances traveled and
proportional distances from the nest within each 4-h
segment as subsamples for each individual. Four-hour
time segments represent reasonable time periods during
which to conduct trapping efforts that coincide with
times of greatest activity away from the burrow. We
determined differences in distance traveled and distance
from the nest among the 4-h time segments by using the
GLM procedure in SAS 9.3. We used Fisher’s protected
least significant difference for mean separation. Due to
difficulty trapping and retaining pocket gophers during
our study, sample size of radio-tagged individuals was
lower than anticipated. Therefore, given the low sample
size, we considered results significant at P,0.10 for all
analyses.
Results
We captured 27 southeastern pocket gophers be-
tween 26 September 2012 and 30 April 2013. One
individual was too small (,100 g) for transmitter
implantation and one adult female (180 g) was in late-
stage gestation. Thus, we implanted transmitters into 25
individuals (mean body mass ¼194 g, SD ¼65, range ¼
122–338). The first 12 individuals captured received
subcutaneously implanted transmitters, four of which
lost transmitters 12–20 d postsurgery. Therefore, we
implanted transmitters in the peritoneal cavity of the
remaining 13 individuals. None of the 13 pocket gophers
with peritoneal implants lost transmitters during the
study, but one died before release due to complications
from surgery. We tracked radio-tagged individuals from 4
October 2012 through 18 September 2013.
Home range
Based on the area-observation curve, a minimum of 17
locations was required to estimate home range. We
recorded the minimum number of locations for 17
individuals. Mean home range size for all pocket gophers
included in the analysis was 921.9 m
2
(SD ¼805.3, range
¼43.4–2246.8; Table S1, Supplemental Material). Home
range size was positively associated with body mass (r¼
0.460, P¼0.063), percent silt at 25 cm (r¼0.554, P¼
0.021), and carbon content at 75 cm (r¼0.457, P¼0.065)
and negatively associated with percent sand at 25 cm (r
¼0.595, P¼0.012), percent clay at 50 cm (r¼0.528, P
¼0.029), and ground cover of grasses other than
wiregrass (r¼0.424, P¼0.089; Table 2; Tables S2–S6,
Supplemental Material).
Survival and cause-specific mortality
All radio-tagged individuals (n¼24; excluding one that
died from surgery complications and was never released)
were used to estimate survival until they were censored
Table 2. Mean (SD), range, correlation coefficient (R), and P
value for variables correlated with home range size of 17 radio-
tagged southeastern pocket gophers Geomys pinetis examined
as part of a study examining home range, survival, and activity
patterns in Baker County, Georgia, 2012–2013. All variables are
percent except pH and body mass (g). Bold variables are
significantly correlated (P,0.10) with home range size.
Variable Mean (SD) Range RPvalue
Body mass
a
180.5 (56.7) 122–338 0.4596 0.0634
Pine straw
a
10.3 (8.6) 0–32 0.0195 0.9407
Leaf litter
b
6.6 (8.5) 0–27 0.1718 0.5098
Woody vegetation
a
5.8 (3.9) 0–13 0.2820 0.2728
Forbs/vines
a
19.9 (11.6) 6–43 0.0525 0.8413
Wiregrass
b
12.1 (20.1) 0–64 0.3622 0.1531
Other grass
a
27.8 (18.3) 4–59 0.4247 0.0893
Sand at 10 cm
b
86.8 (7.8) 57.2–89.6 0.0166 0.9495
Sand at 25 cm
b
86.8 (6.4) 68.8–93.6 0.5948 0.0118
Sand at 50 cm
b
87.4 (7.0) 72.8–93.2 0.2868 0.2643
Sand at 75 cm
b
81.2 (10.2) 60.8–91.6 0.0550 0.8339
Sand at 100 cm
a
79.7 (13.0) 56.4–93.2 0.0174 0.9473
Silt at 10 cm
b
8.6 (3.9) 6.4–22.4 0.1350 0.6055
Silt at 25 cm
b
8.8 (4.6) 4.4–20.8 0.5539 0.0211
Silt at 50 cm
b
6.4 (3.6) 4.8–18.8 0.0835 0.7502
Silt at 75 cm
b
7.6 (4.2) 4.4–20.8 0.2469 0.3395
Silt at 100 cm
b
6.0 (2.2) 4.4–13.2 0.0078 0.9762
Clay at 10 cm
b
4.6 (4.2) 0.8–20.4 0.0654 0.8032
Clay at 25 cm
b
4.4 (2.8) 0.4–14.4 0.3245 0.2038
Clay at 50 cm
b
6.1 (6.2) 0.4–16.4 0.5283 0.0293
Clay at 75 cm
b
11.2 (10.4) 2.0–30.4 0.1266 0.6282
Clay at 100 cm
b
14.3 (11.5) 2.0–30.4 0.1190 0.6491
Nitrogen at 10 cm
b
0.058 (0.020) 0.030–0.123 0.1165 0.6561
Nitrogen at 25 cm
b
0.031 (0.010) 0.000–0.043 0.0504 0.8476
Nitrogen at 50 cm
b
0.022 (0.011) 0.000–0.036 0.1950 0.4534
Nitrogen at 75 cm
b
0.021 (0.011) 0.000–0.033 0.2572 0.3189
Nitrogen at 100 cm
b
0.022 (0.009) 0.000–0.033 0.0308 0.9067
Carbon at 10 cm
a
1.173 (0.591) 0.257–2.615 0.2182 0.4002
Carbon at 25 cm
a
0.435 (0.193) 0.095–0.826 0.0526 0.8412
Carbon at 50 cm
a
0.199 (0.060) 0.116–0.307 0.4019 0.1098
Carbon at 75 cm
b
0.143(0.109) 0.000–0.504 0.4574 0.0649
Carbon at 100 cm
a
0.095(0.047) 0.000–0.205 0.2962 0.2484
pH at 10 cm
b
5.59 (0.60) 5.06–7.29 0.2230 0.3895
pH at 25 cm
a
5.56 (0.58) 4.95–7.35 0.3236 0.2052
pH at 50 cm
a
5.51 (0.54) 4.90–7.32 0.3252 0.2027
pH at 75 cm
a
5.53 (0.38) 5.04–6.57 0.2148 0.4078
pH at 100 cm
a
5.26 (0.54) 4.52–6.69 0.3606 0.1551
a
Relationship determined using Pearson’s product moment correla-
tion.
b
Relationship determined using Spearman’s rank correlation.
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 549
due to transmitter loss/failure (n¼17) or unknown fate
(n¼5; Table S1, Supplemental Material). We documented
two mortality events during the study that are reflected
in the weekly estimates of survival. Survival rate dropped
from 1.000 to 0.857 with a mortality event at week 16
and then dropped to 0.779 with a second mortality event
at week 35, where it remained until the end of the 51-wk
study (Figure 2; Table S7, Supplemental Material). Avian
predation was likely the cause of both mortalities.
Dispersal
Three of the 20 radio-tagged individuals tracked for
.20 days dispersed during the study. All were smaller
(137, 131, and 155 g) than mean body mass of all
individuals (194 g). The first individual dispersed on 25
October 2012 after 22 d of tracking, traveled 264.9 m
over 3 d, and settled at a new location for the remaining
75 d of the tracking period. On 12 November 2012, this
individual made a 305.7-m excursion over 3 d to a third
location, but returned to the second location 3 d later.
The second individual dispersed on 4 November 2012
after 32 d of tracking, traveled 319.1 m over 9 d, and
settled at a new location for the remaining 70 d of the
tracking period. The third individual dispersed on 15
June 2013 after 77 d of tracking, traveled 143.2 m over 10
d, and settled at a second location for the remaining 84 d
of the tracking period.
Daily activity patterns
Mean maximum distance individuals were recorded
from nests during 24 h of focal telemetry was 17.0 m (SD
¼14.1, range ¼3.8–41.9). Mean maximum distance
traveled during 20-min intervals of focal telemetry was
20.5 m (SD ¼18.4, range ¼4.0–51.0). There was no
difference in proportional distance from the nest among
the six 4-h segments (P¼0.139; Figure 3; Table S8,
Supplemental Material). However, proportional distance
traveled differed (P¼0.085) among segments. The mean
separation test indicated that proportional distance
traveled from 1600 to 2000 hours was greater than
proportional distance traveled from 0800 to 1200 hours,
from 1200 to 1600 hours, and from 2000 to 2400 hours,
and proportional distance traveled from 0000 to 0400
hours was greater than proportional distance traveled
from 1200 to 1600 hours and from 2000 to 2400 hours
(Figure 4; Table S9, Supplemental Material).
Discussion
Home range
To our knowledge, this is the first study to report
southeastern pocket gopher home range estimated
using radiotelemetry. Hickman and Brown (1973a)
estimated mean home range size of eight southeastern
pocket gophers in Hillsborough County, Florida, based
on the area in which new mounds were produced. Their
mean estimate (¯
x¼2,666.5 m
2
,SD¼2308.5; calculated
using the rectangular dimensions reported) is almost 3
times larger than our estimate (¯
x¼921.9 m
2
,SD¼805.3).
Hickman and Brown (1973a) delineated ‘‘ maximum
dimensions of mound pattern size’’ that likely included
considerable unused area. In addition, they could not
Figure 2. Estimated survival rate of 24 radio-tagged southeastern pocket gophers Geomys pinetis over a 51-wk study examining
home range, survival, and activity patterns in Baker County, Georgia, 2012–2013. Vertical hash marks indicate censorship events.
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 550
ensure that mounding was by a single individual rather
than two or more in close proximity. In comparison to
other members of Geomyidae, home range of south-
eastern pocket gophers we report is larger than those
reported for the Ozark pocket gopher (¯
x¼291.8 m
2
,SD¼
162.2; Connior and Risch 2010), Botta’s pocket gopher (¯
x
¼474.4 m
2
,SD¼148.2; Bandoli 1987), and the Mazama
pocket gopher Thomomys mazama (¯
x¼108 m
2
,SD¼
Figure 3. Proportional distance of nine radio-tagged southeastern pocket gophers Geomys pinetis from the nest during 4-h time
segments throughout the diel period recorded as part of a study examining home range, survival, and activity patterns in Baker
County, Georgia, 2013. Bars show standard error.
Figure 4. Proportional distance of nine radio-tagged southeastern pocket gophers Geomys pinetis traveled during 4-h time
segments throughout the diel period recorded as part of a study examining home range, survival, and activity patterns in Baker
County, Georgia, 2013. Bars show standard error.
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 551
37.9; Witmer et al. 1996). Pocket gopher home ranges
likely shift as individuals excavate new foraging tunnels,
which could result in an overestimation of home range
size. However, given the time frame in which we tracked
pocket gophers in our study (¯
x¼103 d), we are confident
that any error in home range estimates is negligible.
Similar to the Ozark pocket gopher (Connior and Risch
2010), we found that home range size for the south-
eastern pocket gopher was associated with body mass.
The observed relationship between body mass and
home range size is likely the result of the increase in
metabolism associated with increased body mass
(McNab 1963). As metabolism increases, the area needed
to procure sufficient resources and sustain metabolic
requirements also increases.
Home range size also may be influenced by the ability
to expand and maintain tunnels, which likely explains the
observed relationship between home range and soil
texture. Because southeastern pocket gophers select
sandy soils (McNab 1966; Wilkins 1985, 1987; Simkin and
Michener 2005), it is counterintuitive that home range
size would be negatively associated with percent sand
and positively associated with percent silt. However, in
comparison to silt, sand compacts poorly (Plaster 2013),
making voids in the soil less stable. Thus, an increasing
silt to sand ratio at 25 cm likely increases stability of
tunnels, reducing collapse and allowing pocket gophers
to maintain larger tunnel systems. However, too much
clay in the soil can limit home range size by increasing
the energetic cost of tunnel expansion, explaining the
observed decrease in home range size with increasing
percent clay (Roma ˜
nach et al. 2005). Our results support
those of Warren et al. (in press) who found that
southeastern pocket gophers were more likely to be
present in areas with sandy, loamy sand or sandy-loam
textures throughout the profile relative to areas with
higher clay content and sandy clay loam, loam or clay
loam textures at deeper horizons.
A negative association between home range size and
resource availability is commonly observed in rodents
(Emsens et al. 2013; Lovari et al. 2013) and has been
documented in other geomyi ds (Roma ˜
nach et al. 2005).
Because southeastern pocket gophers feed on above-
and belowground parts of plants (Golley 1962), percent
vegetative ground cover estimated within home ranges
should represent available food resources. Therefore, the
observed negative association between home range size
and ground cover of grasses (other than wiregrass) is
likely due to the ability of pocket gophers to procure
sufficient food in smaller areas when food resource
availability is higher. Southeastern pocket gophers feed
on a variety of herbaceous plants, and grasses are
consumed extensively (Ross 1976). Because foraging
pocket gophers must balance procuring food and
expanding tunnels (Vleck 1981), it would be inefficient
to expand tunnels larger than necessary to gather
sufficient food.
The cause for the positive association between home
range size and soil carbon content at 75 cm is unknown.
Based on our field observations, tunneling was limited at
that depth, and organic matter should not have an
impact on the ability to procure food. Thus, this
observed relationship may be an artifact of a shared
relationship with an unquantified variable.
Survival and cause-specific mortality
The survival rate we observed is similar to the only
comparable study to examine survival in pocket gophers
(Connior and Risch 2010). It is likely that, like most
geomyids, the southeastern pocket gopher has high
survival because its fossorial lifestyle reduces predation
risk. However, predation risk in geomyids is likely higher
during dispersal, which typically occurs above ground
(Vaughn 1963; Williams and Cameron 1984; Daly and
Patton 1990). Although we were unable to document
fate of five individuals, only one of those individuals was
known to disperse. Our observations suggest that
southeastern pocket gophers likely disperse before
reaching 100 g, the minimum weight of individuals
radio-tagged in our study. Therefore, it is unlikely that
the individuals of unknown fate had a significant effect
on our survival estimate. The survival rate observed in
our study supports the contention that longevity in
southeastern pocket gophers is likely more than 2 y
(Brown 1971).
We could not positively identify predators responsible
for the two suspected predation events, but avian
predation was suspected in both cases. In the first
mortality event, we recovered the individual from its
burrow with a puncture wound to the left shoulder and
extensive bruising to the face and muzzle. There was an
opening into the burrow 3–5 m from where the carcass
was recovered, which seemed to be the beginning of a
mound. In the second mortality event, we retrieved the
transmitter near the burrow at the base of a tree. There
were no signs of the carcass. Again, there was an
opening into the burrow 10–15 m from the transmitter,
which seemed to be the beginning of a mound. Pocket
gophers are vulnerable to predation when they emerge
above ground to dispose of soil and repeated trips to the
surface likely attract predator attention (Hickman and
Brown 1973b). We suspect the puncture wounds to the
shoulder and bruising to the muzzle found on the
carcass of the first mortality to be from raptor talons. The
individual apparently escaped the initial attack, but later
died from the injuries. In the second mortality, we
suggest the transmitter was dropped by a raptor feeding
on the carcass. Although snakes in the genus Pituophis
are the primary predators of pocket gophers in regions
where they coexist (Rudolph et al. 2002; Sterner et al.
2002), snakes did not seem to be the cause of either
mortality. The small number of observed mortalities that
occurred during this study likely does not represent the
full range of potential southeastern pocket gopher
predators.
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 552
Dispersal
Interestingly, only 3 of the 20 radio-tracked individuals
in our study dispersed. Because we only radio-tagged
individuals weighing more than 100 g, it is possible that
the individuals we tracked had already dispersed and
established territories at the capture site. Daly and
Patton (1990) also found that dispersal by adult Botta’s
pocket gophers was uncommon. Although one or two
small mounds were present, there were no mound
systems between the initial and final locations of the
dispersing radio-tagged individuals, suggesting that
dispersals occurred above ground. Furthermore, we
incidentally captured four juvenile pocket gophers (89–
95 g) above ground that we assumed to be dispersing.
Aboveground dispersal is common in geomyids (Vaughn
1963; Williams and Cameron 1984; Daly and Patton
1990). Dispersals of three, presumably subadult (155 g)
pocket gophers during our study and the four inciden-
tally captured juveniles that were assumed to be
dispersing suggests that southeastern pocket gopher
dispersal occurs prior to sexual maturity, similar to
Botta’s pocket gopher (Daly and Patton 1990). Juvenile
female Botta’s pocket gophers disperse soon after
weaning and males disperse later as subadults. Unfortu-
nately,wewereunabletodeterminesexofthe
dispersing individuals.
Unlike other studies that documented no evidence of
homing behavior in pocket gophers (Vaughn 1963;
Hickman and Brown 1973a), our observation of a pocket
gopher traveling more than 300 m from its burrow and
returning within 1 wk suggests that the southeastern
pocket gopher may be capable of homing. Homing is a
common ability among rodents and suggests the use of
one of many advanced navigational strategies, such as
memorizing landscape elements (Griffo 1960; Van Vuren
et al. 1997), geomagnetic orientation (August and
Ayvazian 1989), or dead reckoning (Etienne 1992).
Further research is needed to fully evaluate homing
mechanisms used by the southeastern pocket gopher.
Daily activity patterns
Our study is the first to document daily activity
patterns of free-ranging southeastern pocket gophers.
The two metrics used to describe daily activity, distance
traveled and distance from the nest, provided indices of
temporal movement patterns throughout the diel
period. Although we could not confirm the purpose of
the movements, it is reasonable to assume that
movements primarily consisted of foraging, waste
disposal, and reproductive efforts (Baker et al. 2003).
Our observed peak activity periods (0000–0400 and
1600–2000 hours) are in contrast to those of Ross (1980)
who found that captive southeastern pocket gophers
were equally active during all periods. Ross (1980)
acknowledged that activity in captive animals may be
misrepresentative of free-ranging individuals because of
the lack of opportunities for foraging activity. However,
Bandoli (1987) detected a peak in activity between 1500
and 1800 in Botta’s pocket gophers, but did not observe
a second peak as documented in our study.
Implications for translocation
Recognition of longleaf pine communities as floral
biodiversity hot spots (Peet and Allard 1993) and critical
habitat for rare fauna has promoted strong interest in
longleaf restoration (Van Lear et al. 2005). As longleaf
communities are restored, suitable habitat for south-
eastern pocket gophers will become available. Although
our study indicates that southeastern pocket gophers are
capable of dispersing further than previously thought,
the highly fragmented nature of newly available habitats
likely will remain an impediment to natural colonization.
Therefore, translocation will be necessary to reestablish
populations in many situations.
Our results provide important baseline information on
southeastern pocket gophers monitored in situ. Whether
our results are representative of individuals moved to
new locations is unknown. Thus, initial translocation
efforts likely will involve monitoring translocated indi-
viduals via radiotelemetry to assess survival and initial
movements following release. Lessons learned during
our study provide guidance regarding transmitter
implantation. Although subcutaneous transmitter im-
plantation has been used and recommended for radio
tracking other pocket gopher species (Cameron et al.
1988; Connior and Risch 2009b, 2010), we documented
an unacceptably high rate of transmitter loss (33%) on
individuals with subcutaneous implantation. In contrast,
we had no transmitter loss associated with intraperito-
neal implantations. The trade-off is a more invasive
surgery with increased risk of complications. In our study,
one of 13 individuals died (7.7%) due to apparent
surgery-related complications. Zinnel and Tester (1991)
experienced low occurrence of surgery-related mortali-
ties (7.4%) with intraperitoneal implantation in Plains
pocket gophers Geomys bursarius. The reason for the
retention difference of subcutaneously implanted trans-
mitters between our study and others is unknown, but
could be related to species-specific anatomical or
behavioral differences, or to specific surgical techniques.
Although we recommend intraperitoneal implantation
based on our results, we advise consultation with a
veterinarian to determine the most appropriate implan-
tation technique based on study species and specific
surgical techniques.
Beyond improving radiotelemetry methodology, our
results can be applied directly to planning and imple-
menting pocket gopher translocations. First, southeast-
ern pocket gopher activity peaked after dusk, suggesting
that trapping to capture individuals for translocations
may be most effective between 1600 and 2000 hours.
Second, we found that southeastern pocket gophers
disperse before they reach sexual maturity. Adults, like in
other geomyids (Daly and Patton 1990), are generally
sedentary. Therefore, translocating adult individuals may
limit above-ground dispersal movements associated with
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 553
increased predation risk (Vaughn 1963). Third, based on
our estimated home range sizes, southeastern pocket
gophers require approximately 1,000 m
2
per individual,
but soil texture and vegetation may influence home
range size. Thus, an assessment of soil and vegetation
characteristics on translocation sites may be necessary to
ensure adequate spacing between individuals. Finally,
we documented a southeastern pocket gopher homing
more than 300 m. Homing ability can lower site fidelity
and affect success of translocation efforts (Van Vuren et
al. 1997; Villase ˜
nor et al. 2013) and should be considered
in translocation decisions. Although southeastern pocket
gopher translocation distances likely will be much
greater than 300 m, individuals attempting to home
may be at greater predation risk. Fencing or other
barriers may be necessary to prevent above-ground
homing movements during the initial period following
release.
Supplemental Material
Please note: The Journal of Fish and Wildlife Management
is not responsible for the content or functionality of any
supplemental material. Queries should be directed to the
corresponding author for the article.
Table S1. Number of telemetry locations, number of
days tracked, home range size (m
2
), body mass (g),
transmitter fate, and documented dispersal for radio-
tracked southeastern pocket gophers Geomys pinetis
recorded as part of a study examining home range,
survival, and activity patterns in Baker County, Georgia,
2012–2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S1 (32 KB DOCX).
Table S2. pH of soil samples taken in five depth (cm)
increments at the center of southeastern pocket gopher
Geomys pinetis home ranges measured as part of a study
examining home range, survival, and activity patterns in
Baker County, Georgia, 2012–2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S2 (22 KB DOCX).
Table S3. Percent nitrogen of soil samples taken in
five depth (cm) increments at the center of southeast-
ern pocket gopher Geomys pinetis home ranges
measured as part of a study examining home range,
survival, and activity patterns in Baker County, Georgia,
2012–2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S3 (22 KB DOCX).
Table S4. Percent carbon of soil samples taken in
five depth (cm) increments at the center of southeast-
ern pocket gopher Geomys pinetis home ranges
measured as part of a study examining home range,
survival, and activity patterns in Baker County, Georgia,
2012–2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S4 (23 KB DOCX).
Table S5. Percent sand, clay and silt in soil samples
taken in five depth (cm) increments at the center of
southeastern pocket gopher Geomys pinetis home ranges
measured as part of a study examining home range,
survival, and activity patterns in Baker County, Georgia,
2012–2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S5 (49 KB DOCX).
Table S6. Percent ground cover of pine litter,
hardwood leaf litter, woody vegetation, forbs and vines,
wiregrass, and other grass species estimated within five
1-m
2
subplots within 18 m of the center of southeastern
pocket gopher Geomys pinetis home ranges measured as
part of a study examining home range, survival, and
activity patterns in Baker County, Georgia, 2012–2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S6 (35 KB DOCX).
Table S7. Number of at risk individuals, number of
mortalities documented, number of individuals added to
the radio-tagged sample, hazard rate, survival estimate,
and standard error and confidence intervals of estimates
for each week of southeastern pocket gopher Geomys
pinetis survival monitoring during a study examining
home range, survival, and activity patterns in Baker
County, Georgia, 2012–2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S7 (49 KB DOCX).
Table S8. Maximum, mean and mean proportional
distance (m) of southeastern pocket gopher Geomys
pinetis telemetry locations from nests recorded at 20-min
intervals during 4-h time blocks across the 24-h diel
period as part of a study examining home range, survival,
and activity patterns in Baker County, Georgia, 2012–
2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S8 (37 KB DOCX).
Table S9. Mean maximum distance, mean distance
and mean proportional distance traveled (m) by south-
eastern pocket gophers Geomys pinetis recorded at 20-
min intervals during 4-h time blocks across the 24-h diel
period as part of a study examining home range, survival,
and activity patterns in Baker County, Georgia, 2012–
2013.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S9 (37 KB DOCX).
Reference S1. Couch GA, Hopkins EH, Hardy PS. 1996.
Influences of environmental settings on aquatic ecosys-
tems in the Apalachicola-Chattahoochee-Flint River
basin. Atlanta, Georgia: US Geological Survey. Water-
Resources Investigations Report 95-4278.
Found at DOI: http://dx.doi.org/10.3996/032017-
JFWM-023.S10 (10989 KB PDF).
Southeastern Pocket Gopher Home Range, Survival, and Activity A.E. Warren et al.
Journal of Fish and Wildlife Management | www.fwspubs.org December 2017 | Volume 8 | Issue 2 | 554
Acknowledgments
We thank Jean Brock for assistance with ArcMap, Mary
Cobb for assistance preparing soil samples, and
Brandon Crouch for analyzing soil samples. We thank
the Joseph W. Jones Ecological Research Center at
Ichauway and the Warnell School of Forestry and
Natural Resources for funding and logistical support.
We thank the Associate Editor and two anonymous
reviewers for valuable input.
Any use of trade, product, website, or firm names in
this publication is for descriptive purposes only and does
not imply endorsement by the U.S. Government.
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