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Spatial relationships of the desert box turtle (Terrapene ornata luteola) captured at a stock tank in Southeastern Arizona

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I radiotracked Desert Box Turtles (Terrapene ornata luteola) captured at a stock tank in desert grassland habitat in southeastern Arizona from 2001-2003. Individual turtles lived in well-defined 95% MCP home ranges, which included the stock tank and overlapped with home ranges of other turtles. Turtles moved 135 m daily in home ranges averaging 9.8 ha and 469 m across greatest axis. Turtles typically returned to the burrow from which they began their daily movements. There were no sexual differences in daily movement or home range size, nor were there annual differences in daily movement among individuals. Seven of 11 turtles made temporary long distance movements averaging 1,066 m away from their home ranges. Six of 11 turtles exhibited significant directionality in their movements relative to the stock tank. Terrapene o. luteola is a species of conservation concern in Arizona. The spatial information presented in this paper can assist science-based management decisions because it contributes to a better understanding of the core habitat requirements of T. o. luteola.
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Herpetological Conservation and Biology 9(1):156162.
Submitted: 7 December 2013; Accepted: 1 March 2014; Published: 13 July 2014.
156
SPATIAL RELATIONSHIPS OF THE DESERT BOX TURTLE (TERRAPENE
ORNATA LUTEOLA
) CAPTURED AT A STOCK TANK IN SOUTHEASTERN
ARIZONA
MICHAEL V. PLUMMER
Department of Biology, Box 12251, Harding University, Searcy, Arkansas 72143, USA, e-mail: plummer@harding.edu
Abstract.—I radiotracked Desert Box Turtles (Terrapene ornata luteola) captured at a stock tank in desert grassland
habitat in southeastern Arizona from 2001-2003. Individual turtles lived in well-defined 95% MCP home ranges, which
included the stock tank and overlapped with home ranges of other turtles. Turtles moved 135 m daily in home ranges
averaging 9.8 ha and 469 m across greatest axis. Turtles typically returned to the burrow from which they began their
daily movements. There were no sexual differences in daily movement or home range size, nor were there annual
differences in daily movement among individuals. Seven of 11 turtles made temporary long distance movements
averaging 1,066 m away from their home ranges. Six of 11 turtles exhibited significant directionality in their movements
relative to the stock tank. Terrapene o. luteola is a species of conservation concern in Arizona. The spatial information
presented in this paper can assist science-based management decisions because it contributes to a better understanding of
the core habitat requirements of T. o. luteola.
Key Words.—Box Turtle; conservation; core area; directionality; home range; migration; movement; Terrapene ornata luteola
INTRODUCTION
Terrapene ornata is a widely distributed terrestrial
turtle ranging from Indiana to western Arkansas and
Louisiana west to South Dakota, Colorado, southeastern
Arizona, and northern Mexico (Legler 1960; Dodd
2001). The species is commonly recognized to be
comprised of two subspecies (questioned by Martin et al.
2013). In the U.S., the Ornate Box Turtle (T. o. ornata)
is the more widely-distributed form found in the central
plains from South Dakota to southern Texas whereas the
Desert Box Turtle (Terrapene o. luteola) is confined to
the arid desert-grasslands of western and central New
Mexico and southeastern Arizona (Legler 1960; Dodd
2001). Whereas extensive field studies of T. o. ornata
populations in Kansas (Legler 1960; Metcalf and
Metcalf 1970), Texas (Blair 1976), Wisconsin (Doroff
and Keith 1990), Nebraska (Converse et al. 2002), and
Iowa (Bernstein et al. 2007) have been reported over the
last fifty years, field studies of T. o. luteola are limited to
one population in New Mexico (Nieuwolt 1996;
Germano and Nieuwolt-Dacanay 1999) and one
population in Arizona (David Hall and Robert Steidl.
2003, unpubl. report; Plummer 2003, 2004). Because T.
o. luteola is a species of conservation concern in Arizona
(Hall and Steidl, unpubl. report), knowledge of its core
area requirements, including the extent and use of space,
is needed to assist possible management decisions
(Semlitsch and Bodie 2003; Congdon et al. 2011).
Terrapene spp. typically live in well-defined, variable-
sized home ranges to which they attempt to return if
displaced either accidently or after making long distance
movements due to nesting, overwintering, or exploration
(Dodd 2001). The extent to which T. ornata adheres to
this general pattern of space use at its southwestern
range margin is poorly known. The primary goal of this
study is to describe the use of space by individual T. o.
luteola in an apparent declining and poorly known
population in southeastern Arizona.
M
ATERIALS AND METHODS
The study area was active ranchland located in the
semi-arid grasslands of the Sulphur Springs Valley, 44
km SE of Willcox, Cochise County, Arizona at 1,400 m
elevation. Vegetation consisted mainly of grasses and
shrubs. Grasses included Love Grass (Eragrostis), Red
Three-awn Aristida), Tabosa (Pleuraphis), Gramma
(Bouteloua), Vine Mesquite (Panicum), and Sacaton
(Sporobolus). Shrubs included Mesquite (Prosopis),
Burro Weed (Isocoma), Catclaw (Acacia), and Russian
Thistle (Salsola). Other plants include Mormon Tea
(Ephedra), Yucca (Yucca), and Prickly Pear (Opuntia).
Mounds of Bannertail Kangaroo Rats (Dipodomys
spectabilis) were conspicuous landscape features and
mound burrows served as the primary subterranean
retreat for T. o. luteola. Most of the annual activity of T.
o. luteola at the site occurs in a distinct 8–12 wk summer
monsoon beginning in early July in which over 50% of
the 31-cm annual rainfall occurs. However, monsoon
rainfall is far from being uniformly distributed in space
or time. Monsoon showers are typically small in
diameter and move about in an apparent unpredictable
and wandering fashion such that some areas receive
abundant rainfall while nearby areas receive no rain
(Michael Plummer, pers. obs.).
Copyright ©2013. Michael Plummer. All Rights Reserved.
Herpetological Conservation and Biology
157
I initially captured all turtles by hand at an approx.
0.1-ha stock tank designated Turtle Tank (TT; Fig. 1). I
used binoculars to search for turtles visiting TT each day
during the typical 3-h morning surface activity period of
T. o. luteola at this site (0600–0900; Plummer 2003).
After a brief period of observation, I captured and
marked each turtle individually by filing a unique
combination of notches in the marginal scutes. I
attached a Model SM1 (AVM Instrument Company,
Ltd., Colfax, California, USA) radiotransmitter to the
posterior-most marginal scutes of 13 turtles with
stainless steel wire (Plummer 2003). Transmitter masses
were well below the recommended 10% of body mass
for reptiles.
I began to systematically observe and mark turtles at
TT on 1 August 2000. I began attaching transmitters on
19 July 2001 and tracked turtles from 19 July - 9
October 2001 and 6 July - 12 September 2002.
Transmitters remained on individual turtles for the
duration of the study. I captured each turtle at the
beginning of each field season to either replace the
transmitter battery (2001 and 2002) or to remove the
transmitter and release the turtle at the termination of the
study (1 August 2003). While tracking, I made
occasional focal observations through binoculars on the
behavior of turtles during their movements.
I used Biotas™ version 1.03.1a (Ecological Software
Solutions LLC, Hegymagas, Hungary) to plot telemetry
relocations and calculate 95% Minimum Convex
Polygon (MCP) home ranges for turtles with at least 20
telemetry relocations. I also used Biotas™ to measure
the longest axis of home ranges and to identify sojourns
as temporary movements that exceeded the length of the
greatest home range axis beyond the home range
boundary. I used Google Earth
TM
version 7.1.1.1888
(Google Inc., Mountain View, California, USA) to
generate an aerial photograph of the study area, to
determine compass headings, and to measure distances
(meters) on the image with the ruler tool. I checked the
data for normality and equality of variances and used
appropriate parametric (t-tests) and non-parametric
(Mann-Whitney and Wilcoxon) tests with SYSTAT 13
(SYSTAT Software, Inc., Chicago, Illinois, USA). I
used Oriana version 4.02 (Kovach Computing Services,
Pentraeth, Isle of Anglesey, Wales, U.K.) for circular
statistical analysis (Rayleigh test) to determine if
telemetry relocations were directional relative to TT. I
calculated mean compass headings for turtles that
demonstrated significant directionality. Alpha was set at
0.05 for all tests. Descriptive statistics are presented
as mean ± SE.
R
ESULTS
I marked 39 individual turtles that visited TT and
recaptured them a total of 75 times over four years.
Eighteen turtles were observed visiting TT in 2000, 17 in
2001, 12 in 2002, and 17 in 2003. The number of turtles
FIGURE 1. Stock tank in Cochise County, Arizona where turtles were initially collected from 2000-2002. The green ring indicates a higher
previous water level.
Plummer.—Spatial Relationships of Desert Box Turtles in Arizona.
158
seen at TT per day ranged from 1–4. Duration of TT
visits averaged 9 ± 1.5 min (range 2–17 min). Behaviors
at TT consisted of drinking, soaking, and foraging along
the shoreline. Turtles were wary when approaching TT
and quickly retreated if they detected my presence.
A total of 13 turtles were tracked over 1–2 years, but
two transmitters failed early during the course of the
study. As a result, movement analyses were based on
data from 11 turtles (five males; six females) each with
at least 20 telemetry fixes (Table 1). The majority of
turtle movements were in well-defined overlapping 95%
MCP home ranges that were highly variable in size and
averaged 9.8 ± 2.63 ha (range 3.6–34.6 ha; CV = 89%)
in area and 469 ± 49.0 m (range 296–821 m; CV = 35%)
in the longest axis (Table 1). Home ranges of 10 turtles
included TT proper; the 95% MCP home range of the
11
th
turtle was within 150 m of TT (Fig. 2A). Home
range area did not differ between the sexes (U = 23.0, df
= 1, P = 0.140) nor between turtles tracked two years
(11.9 ± 3.98 ha) compared to turtles tracked one year
(6.1 ± 0.91 ha; U = 9.00; df = 1, P = 0.345). Daily
movement averaged 135 ± 13.6 m and did not differ
between years (paired t = 1.52, df = 21, P = 0.18) or
sexes (t = 0.762, df = 9, P = 0.47).
Long-distance movements beyond the turtles’ home
ranges averaged 1,066 m (SD = 142 m, min/max =
431/1,485 m) and were made within a single day by
seven individuals (three males; four females; Table 1).
The average long distance one day sojourn was eight
times the distance of the overall average daily
movement. Sojourn length did not differ between the
sexes (t = 0.50; df = 5, P = 0.64). The path taken by one
female during a sojourn exhibited minimal tortuosity and
often followed portions of cattle paths. Her emigration
and immigration paths were essentially identical. She
gained > 5% in body mass during her sojourn.
The SE quadrant around TT contained the fewest fixes
pooled for all turtles (SE = 9.3%; NE = 34.5%; SW =
38.0%; NW = 18.2%). Six of 11 turtles (three males;
three females) exhibited significant directionality
relative to TT (Table 1) but only one turtle (AKW) had a
mean compass heading in the SE quadrant (Table 1; Fig.
2B). The SE quadrant contained a lower-lying wash,
several human structures, disturbed areas associated with
an old corral, and a junction of primitive dirt roads
within approx. 600 m of TT (Fig. 2C).
D
ISCUSSION
A portion of the home ranges of 10 of 11 radiotracked
turtles captured at TT encompassed TT, whereas the 11
th
turtle maintained a home range near TT. Assuming the
remaining 28 turtles observed at TT but not radiotracked
also maintained home ranges at TT, a minimum of 39
turtles regularly used TT. Because stock tanks are
artificial human structures, the relationship between
turtles and TT is likely a facultative response of turtles
living in a hot, semiarid environment in which most of
the sparse annual rainfall is restricted to a few weeks of
the year.
Several factors could affect home range size at the
study area including the presence or absence of free
water. For example, the 95% MCP home ranges of T. o.
luteola in a nearby study measured 9 ha for turtles
having home ranges within 2 km of a permanent water
source but only 2.8 ha when home ranges were distant
from a permanent water source (Hall and Steidl, unpubl.
report). The small home ranges distant from a water
source are comparable in area to the MCP home ranges
of T. o. luteola in a study located 350 km NE of TT on
the Sevilleta National Wildlife Refuge in New Mexico
(1.6 ha; Nieuwolt 1996). Whether Nieuwolt’s (1996)
study site contained a permanent water source is
unknown but an examination of that study site on
Google Earth (1996 imagery) revealed the nearest stock
tank to be approx. 1,200 m outside the study site
TABLE 1. Movement characteristics of five male and six female Desert Box Turtles (Terrapene ornata luteola) in desert grassland habitat in SE
Arizona and the results of Rayleigh tests of random distribution relative to Turtle Tank (TT) from 2000–2003. Shown are 95% Minimum
Convex Polygon home ranges, maximum length of MCP home ranges, mean daily movement, and temporary movements outside of MCP home
ranges (sojourns). Mean compass headings for turtles showing significant directionality are relative to the center of TT.
ID Sex
No.
Telemetry
Locations
No. Years
tracked
95%
MCP
(Ha)
Max HR
Axis (m)
Mean
Daily
Movement
(m/d)
Sojourn
Length
(m)
Mean
Compass
Heading
(°)
Rayleigh
Test
Z
Prob.
AMN M 66 2 11.8 574 146 1,179 - 1.24 0.289
AJW M 64 2 34.6 821 238 977 308 7.56 < 0.001
AKM M 58 2 8.3 380 105 - - 0.43 0.648
ALU M 46 2 7.4 427 136 781 301 9.05 < 0.001
AKW M 42 1 5.9 368 81 - 159 9.90 < 0.001
AJU F 34 2 5.1 296 192 1,469 7 13.73 < 0.001
ALN F 65 2 3.6 347 105 1,485 - 1.31 0.271
AMV F 21 1 4.3 313 111 431 249 14.53 < 0.001
ALO F 24 1 8.6 668 142 1,141 - 0.84 0.923
CIX F 20 1 5.4 480 129 - 255 18.92 < 0.001
AJN F 70 2 12.8 484 103 - - 1.08 0.340
Herpetological Conservation and Biology
159
boundaries. Another factor that could affect home range
size is habitat degradation, which has been reported to
increase home range size of T. o. ornata in Wisconsin
(Curtin 1997). The Sulphur Springs Valley has been
subjected to long-term drought, suburbanization,
livestock overgrazing, and invasion by exotics
(especially Mesquite), which has resulted in severe
degradation of native desert-grassland habitats in many
areas (Bailey 1994; Rosen et al. 1998). Finally, the
duration of tracking may affect home range size in
turtles. For example, home ranges of Emydoidea
blandingii calculated from data collected over several
years were larger than annual home ranges (Schuler and
Thiel 2008). In this study, I could not detect a difference
in home range size for T. o. luteola based on data
collected over one year vs. those collected over two
F
IGURE 2. A) 95% MCP home ranges of 11 Desert Box Turtles (T. o. luteola) relative to one another and to TT (closed circle in square); B)
Circular histogram showing percentage of compass headings of daily movements for six T. o. luteola (three males; three females) that
demonstrated significant directionality in their movements relative to TT. C) Aerial view of Turtle Tank (TT; dry in this photograph) and
surrounding area. Light lines identify primitive dirt roads.
A
B
TT
Corral
C
Plummer.—Spatial Relationships of Desert Box Turtles in Arizona.
160
years, suggesting that T. o. luteola individuals move
through their entire home range each year.
Alternatively, I may have had insufficient statistical
power to detect an existing difference due to a low
sample size.
Home range sizes in this study should not be
compared to those reported for T. o. luteola in an earlier
study at this site in which data were collected over a
brief period and calculated with a method designed to
yield an index of relative home range sizes rather than
absolute sizes (Plummer 2003).
As reported in an earlier study (Plummer 2003),
surface activity and movements were enhanced by local
rainfall. Movements appeared to be mostly associated
with foraging and were made primarily on bare soil in
sparsely vegetated areas. Although the sinuous paths of
daily movements appeared to have no directional
predisposition (Claussen et al. 1997; Plummer 2003),
turtles were familiar with their home ranges and aware
of their location because they typically returned to the
burrow from which they began their early morning and
late afternoon foraging episodes (Plummer 2003).
While foraging, turtles maintained a high level of
vigilance and quickly responded to disturbances by
ceasing movement and extending their legs and
sometimes their neck vertically in apparent surveillance
behavior (Plummer 2003). Similar behavior, followed
by a quick retreat, was observed at TT when turtles
detected my presence.
Temporary sojourns, commonly reported for T. o.
ornata and its close relative T. carolina, may result from
feeding and nesting forays, exploratory movements,
movements to and from overwintering or nesting sites,
and movements due to environmental perturbations
(Dodd 2001). The most plausible functions for sojourns
in my study were feeding forays and exploratory
movements. Sojourn movements associated with
overwintering and environmental perturbations were
non-factors and the weight gain of one female followed
during her sojourn is inconsistent with a nesting
function.
Differences in habitat, elevation, relief, and vegetation
were difficult to detect around most of TT. However,
the SE quadrant contained a slightly lower wash area
with more dense vegetation. The non-random
distribution of radiotelemetry fixes suggests turtles may
have responded to the slight differences in habitats
around TT. Furthermore, the paucity of turtle fixes in
the SE quadrant, which contained several human-
disturbed areas, suggests avoidance of degraded habitats.
Conservation.—All species and subspecies of
Terrapene are of national concern in the U.S. and are
listed in CITES Appendix II. Terrapene o. luteola is
listed as imperiled (S2) or vulnerable (S3) in Arizona
(NatureServe. 2013. NatureServe Explorer: An Online
Encyclopedia of Life. Available from
http://www.natureserve.org/explorer
. [Accessed: 31
October 2013]). In a status assessment study in Arizona,
Hall and Steidl (unpubl. report) reported that T. o.
luteola may be locally common but has declined
statewide. Together these results suggest that T. o.
luteola may need to be considered for conservation
management to assure future population viability. The
results reported herein on home range size, home range
fidelity, internal home range movements, and temporary
movements out of the home range should be of value to
science-based conservation efforts, especially to
questions of how individual turtles use their habitat and
how turtles might respond when involuntarily removed
from their home ranges in repatriation, translocation, and
relocation efforts (Dodd 2001; Hill et al. 2009, Refsnider
et al. 2012). Nonetheless, additional biologically
relevant information is still needed for a more complete
understanding of the core area requirements of T. o.
luteola. For example, data are needed on potential
nesting migrations out of the home range and on the area
necessary to buffer core habitat from the edge effects of
surrounding land use (Semlitsch and Bodie 2003;
Congdon et al. 2011).
Although individual T. o. luteola are clearly attracted
to stock tanks, whether the attraction affects turtle
behavior, physiology, and life history is unknown.
Historically, water gained from foraging and drinking at
monsoon rain pools provided sufficient water for
survival and population growth; water available at stock
tanks is the result of relatively recent human activities.
Water supplementation is known to increase surface
activity and food acquisition in the long-lived desert
dwelling Gila Monster (Heloderma suspectum; Davis
and DeNardo 2009) and increase growth in the Desert
Tortoise (Gopherus agassizii; Field et al. 2007).
Providing additional aquatic resources might similarly
benefit individual turtles; however, there are no
comparative data to support or refute these ideas.
The distribution of stock tanks could be an influential
force shaping the metapopulation dynamics of T. o.
luteola (Roe and Georges 2007; Roe et al. 2009) if stock
tanks represent clumped resources for turtles and local
population density is increased as a result. An
understanding of these issues would require comparative
population studies in areas with and without stock tanks.
Developing comprehensive management strategies for T.
o. luteola on ranchlands will require not only spatial
information such as provided in this paper, but also an
understanding of turtle population structure and how
stock tanks might affect that structure.
Acknowledgments.—Jho and Valer Austin allowed me
to work and reside on their ranchland. Jim and Peg Tout
and Tony and Helen Fraze provided numerous courtesies
and willingness to share their observations on box
Herpetological Conservation and Biology
161
turtles. Chance Fraze helped collect turtles. Justin
Congdon and Dick van Loben Sels provided ideas,
supplies, use of equipment, and assistance of various
kinds. Julie Fuller and Amanda Moors assisted in
radiotracking. Bob Steidl and David Hall provided their
unpublished T. o. luteola report to the Arizona Game and
Fish Department Heritage Program. Howard Snell
provided information on Nieuwolt’s (1996) study site in
New Mexico. Justin Congdon and Nathan Mills
commented on the manuscript. Authorization to collect
and monitor turtles was provided by the Arizona Game
and Fish Commission (Scientific Collecting Permits
#SP772582, #SP834284). Methods of study followed
established guidelines for reptilian field research
(http://www.asih.org/sites/default/files/documents/Resou
rces/guidelinesherpsresearch2004.pdf). This research
was funded by several Faculty Development Grants from
Harding University.
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M
IKE PLUMMER is professor and two-time former Chairman of the Department of Biology at Harding University
where he teaches biostatistics and herpetology. He holds a Ph.D. from the University of Kansas and a M.S. from
Utah State University. His research focuses on the ecology of snakes and turtles on which he has published for 40
years. (Photographed by Chance Fraze).
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