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2011 SOUTHEASTERN NATURALIST 10(1):39–52
Investigation of Adult Male White-tailed Deer Excursions
Outside Their Home Range
Gabriel R. Karns1,2,*, Richard A. Lancia1, Christopher S. DePerno1,
and Mark. C. Conner3
Abstract - Although male and female Odocoileus virginianus (White-tailed Deer) exhibit
high site fidelity throughout the year, individuals occasionally leave their home ranges on
short excursions during the fall and winter months. Although motives for these extraneous
movements are difficult to discern, excursions are likely the function of the breeding
season, food sources, limited escape cover, and/or human disturbances. From 2003–2007,
we examined GPS collar locations of 32 adult male White-tailed Deer at Chesapeake
Farms, MD. Seasonal excursions (n = 37), defined as movements lasting a minimum
of 6 hours and venturing at least 0.5 km from 95% kernel home-range contours, were
examined relative to possible motives related to food resources, breeding, and hunting
pressure. Sixty-three percent (n = 20) of adult males made at least one excursion outside
their home range immediately before or during breeding season. Based on the seasonal
timing of excursions, breeding-season-related motives were likely the driving force behind
the majority of adult male White-tailed Deer excursions, whereas hunting pressure
and food resources were not a probable cause.
GPS data from previous research conducted at Chesapeake Farms, MD,
indicated that some male Odocoileus virginianus (White-tailed Deer) traveled
significant distances outside their normal home range for periods lasting between
6–28 hours, with most excursions occurring immediately before and during the
fall breeding season (Tomberlin 2007). Just before breeding season, male activities
intensify (i.e., rubbing, scraping, sparring, and searching for estrous females)
and movement and home ranges increase (Guyse 1978, Hawkins and Klimstra
1970, Hosey 1980, Tomberlin 2007). Additionally, White-tailed Deer may temporarily
leave their home range to avoid hunting pressure and other disturbances
(Hood and Inglis 1974, Naugle et al. 1997, Vercauteren and Hygnstrom 1998).
Dispersal movements are predominantly made by juvenile (1.5-year-old) male
White-tailed Deer and result in permanent emigration (Brinkman et al. 2005,
McCoy et al. 2005, Rosenberry et al. 1999, Shaw 2005), whereas excursions are
temporary movements outside an established home range.
As estrus approaches, females concentrate movement and scent markings
within their core areas (Fraser 1968, Holzenbein and Schwede 1989, Ivey and
1Fisheries and Wildlife Sciences Program, Department of Forestry and Environmental
Resources, North Carolina State University, Box 7646, Raleigh, NC 27607. 2Current address
- School of Forestry and Wildlife Sciences, 3301 Forestry and Wildlife Sciences
Building, Auburn University, Auburn, AL 36849-5418. 3DuPont Crop Protection, Chesapeake
Farms, 7319 Remington Drive, Chestertown, MD 21620. *Corresponding author
40 Southeastern Naturalist Vol. 10, No. 1
Causey 1981, Marchinton 1968, Nelson and Mech 1981), which may increase
the chance of males detecting females by focusing activities within a small area
(Holzenbein and Schwede 1989, Ozoga and Verme 1975). As females enter estrus,
males stimulated by olfactory and behavioral cues instigate male-male competition
(Cox and Boeuf 1977) and separate individual females from the matriarchal
herd and tenaciously follow them for 1–6 days (Brown 1971, Crawford 1962,
Hawkins and Klimstra 1970, Holzenbein and Schwede 1989). By luring courting
males into a chase and venturing outside her core area, females might attract attention
from other potential mates. Once engaged in the chase, males might easily
be led outside their home range and into unfamiliar territory, possibly bringing
multiple males together and stimulating intrasexual competition (Cox and Boeuf
1977, Emlen and Oring 1977). After being tended and bred, females will decrease
activity, return to core areas, and resume normal levels of movement and activity
(Cox and Boeuf 1977, Holzenbein and Schwede 1989, Ozoga and Verme 1975).
However, females not bred might initiate a search strategy to find a mate during
her 24-hour window of receptivity (Ozoga and Verme 1975). Excessive movement
of females during the rut might indicate poor male breeding performance
in a herd (Holzenbein and Schwede 1989) or may be a byproduct of extremely
low deer densities where the odds of two deer encountering one another are small
(D’Angelo et al. 2004). In rare instances, females may make excursions outside
their home range during the breeding season even with abundant mature males in
the population (Kolodzinski 2008).
Excursions by adult male White-tailed Deer are not easily studied because
of their unpredictability and difficulty of detection using conventional radio telemetry
equipment. Excursions may represent exploratory searches for estrous
females (Guyse 1978, Hawkins and Klimstra 1970, Hosey 1980, Moore and
Marchinton 1974), a new food source, or may be a male chasing an unreceptive
female (Richardson and Petersen 1974), a male being led by an estrous female
back to her core area (Cox and Boeuf 1977, Holzenbein and Schwede 1989),
a male leading a receptive mate away from intrasexual breeding competition
(Moore and Marchinton 1974), or female incitation of male competition (Cox
and Boeuf 1977). However, excursions by adult males are not fully understood
and may not be limited to these motives. Hence, our objectives were to describe
adult male White-tailed Deer excursions during the fall and winter of 2003–2007
at Chesapeake Farms and compare male and female movements to identify simultaneous
We conducted the research at Chesapeake Farms located on the Eastern Shore
of the Chesapeake Bay in Kent County, MD, 10 km southwest of Chestertown
(39°10'N, 76°10'W), with a mean elevation of 13 m above sea level (McLeod
and Gates 1998). Owned and operated by DuPont Agricultural Enterprise,
Chesapeake Farms was a 1300-ha wildlife management and agricultural research
demonstration area. Approximately 50% of the study area was forested with nonalluvial
swamps that consisted primarily of Quercus spp. (Oaks), Liquidambar
2011 G.R. Karns, R.A. Lancia, C.S. DePerno, and M.C. Conner 41
styraciflua L. (Sweetgum), Nyssa sylvatica Marsh. (Black Gum), and Acer
rubrum L. (Red Maple). Smilax spp. (Greenbriar), Clethra alnifolia L. (Sweetpepper
Bush), and Vaccinium corymbosum L. (Highbush Blueberry) dominated
the understory. Cash crops (Zea mays L. [Field Corn], Glycine max (L.) Merr.
[Soybeans], and Triticum spp. [Winter Wheat]) composed 20% of the study area.
Fallow fields composed 13% of the farm (Dactylis glomerata L. [Orchard Grass],
Trifolium spp. [Clover], Sorghum spp. [Sorghum], and Lolium multiflorum Lam.
[Rye]). The remaining 17% was composed of nonforested wildlife cover and
manmade waterfowl impoundments (Shaw 2005).
Shaw (2005) estimated deer density to be 33 deer/km2, and the majority of
deer harvest occurred during Maryland’s two-week shotgun season from the
first Saturday after Thanksgiving for two continuous weeks. Because of initial
antler-harvest restrictions (≥7 points) implemented in 1994 and more stringent
criteria (≥40 cm outside antler width) enforced in 1997, male age class structure
on Chesapeake Farms shifted from a traditional harvested population (mostly
1.5-year-old males) to predominantly ≥2.5-year-old males (Shaw 2005). The fall
2007 male:female ratio estimate was 1:1.5 and has remained biologically stable
since 2002, when intensive doe harvest to curb excessive crop depredation ceased
(M. Conner, Chesapeake Farms, Chestertown, MD, unpubl. data).
From June–August 2003–2007, we captured 32 (2003 [n = 3], 2004 [n = 3],
2005 [n = 9], 2006 [n = 10], and 2007 [n = 7]) adult male (≥2.5 years old) Whitetailed
Deer. In the field, we estimated deer age using antler and body characteristics
(Richards and Brothers 2003). We used a Dan-Inject JM Standard dart projector
(Dan-Inject, Inc., Fort Collins, CO) and 3-ml radio transmitter darts (Pneu-dart,
Inc., Williamsport, PA) to administer anesthetic drug combinations of 2.4 ml Telazol
(200 mg/ml; Fort Dodge Animal Health, Fort Dodge, IA) and 0.6 ml Xylazine
(450 mg/ml; Wildlife Laboratories, Inc., Fort Collins, CO) (Kreeger et al. 2002) or
0.5 ml Medetomidine (20 mg/ml; Wildlife Laboratories, Inc., Fort Collins, CO),
1.0 ml Ketamine (200 mg/ml; Fort Dodge Animal Health, Fort Dodge, IA) and 1.4
ml Telazol (200 mg/ml) (Muller et al. 2007). The transmitter dart allowed us to
track the deer using radio telemetry equipment (Telonics, Inc., Mesa, AZ). If the
animal was not fully sedated when located, we administered an additional 1.1 ml
Ketamine (200 mg/ml) booster intramuscularly.
Once fully immobilized, we applied eye ointment (Paralube, Pharmaderm,
Melville, NY) to prevent corneal drying and blindfolded the animal to minimize
stress. The deer were positioned sternally or on their right side for processing. We
monitored vital signs (open airway, pulse, respiration, temperature) initially and
every 15 minutes throughout the procedure. We surgically removed darts, flushed
the puncture with Betadine (Purdue Pharma, L.P., Stamford, CT), and applied
antibiotic cream (Farnam Companies, Inc., Phoenix, AZ) to the wound. A broadspectrum
antibiotic (i.e., LA 200; 1 ml/11.34 kg; Pfizer Animal Health, Exton,
PA) was administered intramuscularly to two sites in the hindquarter. We fitted
each deer with a Lotek 3300L GPS collar (Lotek Engineering, Newmarket, ON,
42 Southeastern Naturalist Vol. 10, No. 1
Canada) and tightened them within 8 cm of the neck to accommodate neck swelling
associated with breeding season. Although each collar was equipped with a
32-week time-delay release mechanism, a remote release mechanism allowed us
to disengage the collar in case of emergency. To assist in field identification, each
deer received colored and numbered cattle ear tags (National Band and Tag, Co.,
Newport, KY). Also, we placed uniquely numbered Monel tags (National Band
and Tag, Co., Newport, KY) in both ears.
At 70 minutes post injection, we reversed Xylazine/Telazol anesthetized deer
with 3.3 ml Tolazoline (100 mg/ml; Lloyd Laboratories, Shenandoah, IA), half
intramuscularly and half intravenously. We used 10.0 ml atipamezole (5 mg/ml;
Pfizer Animal Health, Exton, PA) to intramuscularly reverse deer anesthetized
with Medetomidine/Ketamine/Telazol. Seventy minutes was adequate time for
the Telazol and/or Ketamine to dissipate from the deer and eliminate the risk of
anesthetic relapse (Tomberlin 2007). We monitored deer until they were capable
of independently leaving the processing site. The research protocol was reviewed
and approved by the Institutional Animal Care and Use Committee at North
Carolina State University (#05-024-0).
We programmed 3300L Lotek GPS collars to collect hourly fixes from June–
February 2003–2007, 20 minute fixes from 5 November–12 December 2006,
and 5-minute fixes from 15 October–15 December 2007. GPS collars recorded
geographic coordinates, date, time, environmental temperature, fix status, and a
position dilution of precision (PDOP) value with each fix. Collars were equipped
with a mortality sensor that triggered after 8 hours of inactivity and emitted a double
pulse VHF signal. To ensure collars were properly functioning and study animals
were alive, deer were monitored twice weekly using radio telemetry equipment.
To delete possible erroneous fixes from the dataset, we filtered fixes through
a pre-determined set of quality-control screenings and omitted all three dimensional
(3D) fixes with PDOP > 10 and two dimensional (2D) fixes with PDOP
> 5 from analyses (Adams 2003, D’Eon and Delparte 2005, Tomberlin 2007). In
addition to the PDOP filter, we omitted all fixes with altitudes outside the range
of -100 m to 100 m (D’Eon and Delparte 2005, Tomberlin 2007). Also, we removed
malfunctioned fixes as indicated by VHF pulse rates or absent VHF signal
After data censoring, we imported GPS fixes for each deer into ArcMap 9.2
(Environmental Systems Research Institute, Inc., Redlands, CA). We projected
all data in Universal Transverse Mercator (UTM) North American Datum (NAD)
1983 Zone 18 North (m). We used top-hour fixes from 7 days post capture to the
end of the data-collection period to generate fixed kernel home ranges (95%)
using Kernel Density Estimator and Percent Volume Contour in Hawth’s Analysis
Tools (Beyer 2004). We chose a smoothing parameter (200) based on close
examination of a wide range of possible values and comparing corresponding
polygons to true distribution of GPS fixes (Laver 2005).
We documented excursions outside of the fixed kernel home ranges (95% volume)
and required movements to exceed 0.5 km from the home-range contour,
encompass six or more continuous hours, and occur between 24 September and
2011 G.R. Karns, R.A. Lancia, C.S. DePerno, and M.C. Conner 43
the collar release date (individual specific). Each movement was classified as
either exhibiting continuous movement throughout the excursion or including a
period of little to no movement (≥3 hours) during the excursion outside the animal’s
home range. Repeated excursions by the same individual were noted and
analyzed independently. We omitted adult male White-tailed Deer movements
transiting between disjunct home ranges.
We defined 24 September–14 October as fall, 15 October–4 November
as prebreed, 5 November–25 November as breeding, 26 November–16 December
as postbreed, and 17 December–collar release date as winter. During
2003–2007, the Maryland two-week firearms season was conducted from
29 November–13 December, 27 November–11 December, 26 November–10
December, 25 November–9 December, and 24 November–8 December, respectively
(Tomberlin 2007). Specific motives for adult male White-tailed
Deer excursions included food-resource explorations (fall, postbreed, and
winter), searching for receptive females (prebreed and breeding), chasing
females (prebreed and breeding), breeding estrous females (breeding), or
hunting-avoidance movements (firearms season).
During May–August 2006, 14 female White-tailed Deer (>1.5 year old) were
captured (same protocol as male White-tailed Deer) with 1.6 ml Telazol (200 mg/
ml) and 1.4 ml Xylazine (200 mg/ml) (reversal: 4 ml of 100 mg/ml Tolazoline)
and collared with Televilt Tellus Basic GPS collars (Televilt/TVP Positioning
AB, Lindesburg, Sweden) to study movement during the breeding and fawning
seasons (Kolodzinski 2008). GPS collars were programmed to collect 45-minute-
interval (1 October–31 January and 1 April–31 July) and 1-hour-interval
(1 February–31 March and August 1–September 30) fixes for 365 days following
deployment (Kolodzinski 2008). To further investigate adult male excursions
during the White-tailed Deer breeding season, we compared female movements
to adult male excursions during the prebreed and breeding periods by overlaying
GPS fixes in ArcMap 9.2.
Thirty-seven adult male White-tailed Deer excursions were documented, with
the highest number (n = 14) occurring during the breed period (Fig. 1). Mean
minimum distance traveled was 778 m (SE = 293 m), with a range of 506–1500 m
(median = 665 m). Mean excursion duration was 10.5 hours (SE = 6.2 hours), with
a range of 6–40 hours. Fifty-nine percent (n = 22) of excursions occurred during the
combined prebreed and breeding periods. Although the number of study animals
steadily decreased because of mortality or collar malfunction, 63% (n = 20) of the
adult males made at least one excursion (Fig. 2). Interestingly, 41% (n = 15) of excursions
were characterized by continuous movement (Fig. 3), and 59% (n = 22) by
periods of little to no movement (Fig. 4). Also, 59% (n = 22) of excursions occurred
at least partially during daylight hours. During Maryland’s two-week firearms
season, we documented five excursions (2003 [n = 1], 2004 [n = 1], 2005 [n = 2],
and 2007 [n = 1]). On three occasions (during prebreed period [n = 1] and breeding
period [n = 2]), males made repeat excursions to the same locations. None of the 37
44 Southeastern Naturalist Vol. 10, No. 1
adult male excursions followed the same movement paths at the same time as any
of the collared female White-tailed Deer.
A total of 63% of the GPS-collared adult male White-tailed Deer on Chesapeake
Farms made excursions outside of their home range. Because the majority
Figure 1. Percent of adult male White-tailed Deer excursions by breeding period, Chesapeake
Farms, MD, 2003–2007.
Figure 2. Percent of adult male White-tailed Deer making 0, 1, 2, or 3+ excursions,
Chesapeake Farms, MD, 2003–2007.
2011 G.R. Karns, R.A. Lancia, C.S. DePerno, and M.C. Conner 45
of excursions occurred during the prebreed and breeding periods, breeding season
activities were likely the most common cause of these movements. Because
the deer population at Chesapeake Farms was characterized by older age class
Figure 3. Continuous excursion by a 3.5-year-old adult male White-tailed Deer, Chesapeake
Farms, MD, 8–9 November 2006 (22:00–6:00); 95% fixed kernel home range is
represented by outlined polygon.
46 Southeastern Naturalist Vol. 10, No. 1
males and a nearly balanced sex ratio, the secondary rut (when unbred females
enter their second estrus cycle) was probably insignificant compared to other regions
(Clutton-Brock et al. 1997, Geist 1971). However, a minimal secondary rut
Figure 4. Excursion with a delay by a 3.5-year-old adult male White-tailed Deer, Chesapeake
Farms, MD, 12–13 November 2007 (21:00–3:30); 95% fixed kernel home range
represented by outlined polygon.
2011 G.R. Karns, R.A. Lancia, C.S. DePerno, and M.C. Conner 47
within the core of Chesapeake Farms did not preclude surrounding properties under
different management schemes and resulting herd demographics (i.e., heavily
skewed sex ratio and fewer adult males) from having pronounced secondary rut
activity. Therefore, after the principal breeding period, it is plausible that males
took advantage of later breeding opportunities by venturing outside Chesapeake
Farms’ boundaries during the postbreed period.
Excursions characterized by continuous movement or associated with a period
of no movement indicate that some motives are more likely than others. For example,
a male searching unsuccessfully for receptive females outside his home
range would likely continue moving through previously unoccupied territory (at
least during the time when the deer was wearing the GPS collar) before returning
to his normal home range. However, a male tending a receptive female might
attempt to isolate her from intraspecific competition and mate with her as many
times as possible (Hirth 1977). It is purely speculative, but repeated excursions
by the same male might suggest revisiting a female group to check for receptive
mates, utilizing a food resource that was discovered by previous experiences,
or returning to a natal range. With only three repeat excursions documented, it
would appear that most excursions were not explorations for new food sources as
one or two trips to a distant food source probably would not justify the risk and
energy expenditure of those excursions—making late breeding season activities
or natal range re-visitation more likely explanations.
Five male excursions did occur during Maryland’s two-week firearms season
(all 5 of these excursions occurred within the postbreed study period); however,
it is unlikely they were prompted by hunting pressure. Based on fine-scale
movement data collected during the 2006 (20-minute fixed interval) and 2007
(5-minute fixed interval) firearms season, we compared adult male movement
during daylight hours to known hunter locations (Karns 2008). Whether disturbed
by hunters or vehicles, flight distances never exceeded 600 m, and no deer left
its home range in response to hunting-related disturbances (Karns 2008). Other
researchers (Altmann 1958, Hood and Inglis 1974, Lagory 1987, Naugle et al.
1997) noted some White-tailed Deer temporarily left their home ranges in direct
response to hunting and other intrusive activities, especially where security and
escape cover were limiting factors in a landscape. At Chesapeake Farms, based
on low levels of hunting pressure, abundant cover, and flight distance data from
adult males during 2006 and 2007, we concluded that hunting pressure was an
unlikely explanation for adult male excursions.
Yearling males are disproportionately vulnerable during lengthy dispersal
movements through unfamiliar territory (Nelson and Mech 1986, Roseberry
and Klimstra 1974). Similarly, adult males venturing on excursions are highly
vulnerable to hunter harvest, antagonistic encounters with competing males,
and other mortality factors in areas where they are less intimately familiar
(Swenson 1982). In 2005, a 5.5-year-old study animal was killed in a vehicle
collision while crossing a two-lane road that was not previously encountered
during its collared period; in 2006, a 3.5-year-old research subject was superficially
wounded by an archery hunter while making an excursion. A majority
48 Southeastern Naturalist Vol. 10, No. 1
of excursions occurred at least partially during daylight and exposed deer to
potential hunter harvest.
Although Kolodzinski et al. (2010) used different criteria to quantify excursions
in female White-tailed Deer at Chesapeake Farms, all five females (≥1.5
year old) with complete datasets exhibited at least one extraneous movement
outside 95% home-range contours. Interestingly, only one (made by the 1.5-yearold
female in the study) of the excursions was documented outside the peak of
breeding-season activity (Kolodzinski et al. 2010). Consistent with the findings
of Kolodzinski et al. (2010), D’Angelo et al. (2004) documented female Whitetailed
Deer excursions occurring in close proximity with their estrous cycles. It
is also known that 1.5-year-old females and even 6-month-old female fawns may
enter estrus up to several months later than adult females (Nixon 1971, Ozoga
and Verme 1982). It is likely that the later excursion by the 1.5-year-old female
corresponded with her belated estrus cycle (Kolodzinski et al. 2010). Although
no data is available for the proportion of young breeding females at Chesapeake
Farms, combining the relatively mild climate with abundant resources, it is not
unlikely that at least a small fraction of the female fawns and certainly some
1.5-year-old females are successfully bred during December and subsequent
months as individuals reach puberty. Young females exhibiting late estrous
cycles may account for an extended breeding season and may partially explain
male White-tailed Deer excursions during the postbreed and winter periods as
they pursue additional opportunities to reproduce.
Female excursions might be expected in a low-density population (D’Angelo
et al. 2004) or where sexually mature males were lacking, but Chesapeake Farms
possesses a high-density population and abundant older-aged males. Kolodzinski
et al. (2010) postulated that female White-tailed Deer may be engaging in a discrete
form of mate selection, even in an environment with seemingly abundant
quality mates. With adult males and adult females displaying extraneous movements
during the breeding season, both sexes may be instrumental in increasing
genetic flow and heterozygosity. It is commonly accepted that juvenile male dispersal
is the predominant source of landscape genetic flow in White-tailed Deer
populations (Nelson 1993).
Ideally, to study potential simultaneous movements of males and females, all
deer in the same geographic area would have GPS transmitters. Obviously the
costs and logistics of doing this are nearly prohibitive, especially in a high-density
population like at Chesapeake Farms. Although none of the female movement
data mirrored any adult male excursions during the 2006 prebreed and breeding
periods, we believe this could simply be an artifact of having too few deer with
GPS collars. Studying a lower-density herd might be more feasible logistically,
but the motives for excursions would likely be different. Additional studies are
needed to determine the cause and frequency of extraterritorial excursions of
adult males during breeding and hunting seasons.
Although two adult males used multiple home ranges or exhibited evidence
of adult dispersal, movements in transit between home ranges were
not included in our analyses. Webb et al. (2007) reported that 15% of adult
2011 G.R. Karns, R.A. Lancia, C.S. DePerno, and M.C. Conner 49
male White-tailed Deer (usually 2.5–3.5-year-olds) disperse and permanently
vacate their natal range. We were unable to find compelling evidence in the
literature suggesting reasons why adult males that occupy high quality habitat
(i.e., Chesapeake Farms, MD) would maintain two separate home ranges, and
the behavior may simply be explained as the idiosyncrasies of a few individuals
(less than 10%). In more northern latitudes, where White-tailed Deer populations
migrate between summer and winter ranges, special caution must be taken to
avoid misclassifying migratory movements as extraneous excursions (Brinkman
et al. 2005, DePerno 1998, Nixon et al. 2008, Tierson et al. 1985). Also,
the inherent nature of a fixed kernel home range dictates that some locations
are located outside the 95% contour. Therefore, rigorous criteria (i.e.,
minimum distance and length-of-time requirements) must be selected for
identifying true excursions to keep superficial wandering movements and erroneous
locations from being misclassified as excursions.
It appears the White-tailed Deer breeding season (whether the peak of adult
female conception in mid-November or younger females entering estrus later in
December and January) motivates the majority of adult male excursions, but it is
difficult to hypothesize plausible motives for excursions occurring during the fall
period. At Chesapeake Farms, excursions put individual deer at risk of mortality
(particularly due to hunter harvests and vehicle collisions), but movements into
unfamiliar environments do not occur frequently enough to drastically impact the
survival rate of older age class cohorts. Researchers recognize that yearling male
dispersal plays a major role in landscape ecology processes such as gene flow
and disease transmission (McCoy et al. 2005, Rosenberry et al. 1999, Schauber et
al. 2007); however, the role of adult male White-tailed Deer excursions in these
same processes is commonly overlooked.
We thank DuPont Crop Protection, the Department of Forestry and Environmental
Resources, and the Fisheries and Wildlife Sciences Program at North Carolina State
University for funding this research project. Also, we are grateful to J. Kolodzinski for
allowing us to compare overlapping movements using his data. Thanks to L. Muller at
University of Tennessee-Knoxville for providing darting equipment. Additional thanks to
R. Fleegle, B. Paugh, and J. Tomberlin for field assistance.
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