Effects of Time of Day and Activity Status on Bobcat
(Lynx rufus) Cover-type Selection in Southwestern Georgia
Jordona D. Kirby, Jessica C. Rutledge, Ivy G. Jones, L. Mike Conner, and Robert J. Warren
Southeastern Naturalist, Volume 9, Issue 2 (2010): 317–326
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2010 SOUTHEASTERN NATURALIST 9(2):317–326
Effects of Time of Day and Activity Status on Bobcat
(Lynx rufus) Cover-type Selection in Southwestern Georgia
Jordona D. Kirby1,3, Jessica C. Rutledge1, Ivy G. Jones1,4,
L. Mike Conner1,*, and Robert J. Warren2
Abstract - Lynx rufus (Bobcat) cover-type selection and activity patterns have been
studied in a variety of landscapes in the southeastern United States, but effects of
individual activity status (active or inactive) and time of day (day, night, crepuscular)
on cover-type selection have not been investigated for this species. Therefore, we
investigated Bobcat habitat use in a Pinus palustris (Longleaf Pine) forest in southwestern
Georgia to determine whether activity status of individuals or time of day
affected seasonal cover-type selection. We monitored 43 radiocollared Bobcats from
2001–2004 and determined habitat use at Johnson’s third order of selection (i.e.,
habitat selection within the home range) using Euclidean distance-based analysis.
Bobcats selected (Λ = 0.017, P = 0.001) habitat within their home ranges; however,
although Bobcats are typically classified as crepuscular, neither activity status (Λ =
0.990, P = 0.981) nor time-of-day (Λ = 0.972, P = 0.647) affected cover-type selection.
Bobcats on our study site preferred agricultural areas and other early to midsuccessional
habitats, probably because they produced abundant prey.
Prey abundance and distribution are important determinants of Lynx
rufus Schreber (Bobcat) habitat use throughout their distributional range
(Anderson 1987, Pollack 1951, Rucker et al. 1989). Previous research illustrated
the importance of prey availability on Bobcat cover-type selection in
the Southeast (Godbois et al. 2004). Godbois et al. (2004) found that Bobcats
were located closer to areas in which supplemental food had been provided
for Colinus virginianus Catesby (Northern Bobwhite; hereafter quail) as a
result of the supplemental food locally concentrating rodents that were attracted
to the grains used to feed quail.
Bobcat habitat use is also influenced by availability of resting and denning
sites, protection from environmental extremes, dense cover for hunting
and escape, and freedom from disturbance (Anderson 1987, Bailey 1974,
Boyle and Fendley 1987, Kitchings and Story 1984, Pollack 1951, Young
1958). Although Bobcats are typically classified as nocturnal or crepuscular,
factors potentially influential on cover-type selection, such as activity status
(i.e., whether an animal is active or inactive) and time of day (i.e., whether
1Joseph W. Jones Ecological Research Center, Route 2, Box 2324, Newton, GA
39870. 2Daniel B. Warnell School of Forestry and Natural Resources, University of
Georgia, Athens, GA 30606. 3Current Address - USDA, APHIS, Wildlife Services,
4708 Western Avenue, Suite A, Knoxville, TN 37921. 4Current Address - Kentucky
Department of Fish and Wildlife Resources, 1 Sportsman’s Lane, Frankfort, KY
40601. *Corresponding author - firstname.lastname@example.org.
318 Southeastern Naturalist Vol. 9, No. 2
it is day, night, or a crepuscular time period) have not been studied. When
monitoring wildlife using radiotelemetry, it is critical to obtain locations
such that the data adequately represent the variation in behaviors for the
species of interest (Samuel and Fuller 1994). Commonly, the ideal schedule
for wildlife telemetry data collection includes both diurnal and nocturnal
or 24-hour sampling (Beyer and Haufler 1994, Smith et al. 1981). When
logistical difficulties necessitate deviation from this ideal schedule for data
collection, research results may be questioned (Beier et al. 2006). Thus, we
assessed whether Bobcat cover-type selection varied with activity status or
time of day.
We predicted that active Bobcats would be located more than expected
in association with early successional cover and edges, as we assumed that
most Bobcat activity was associated with their foraging behavior. Similarly,
we predicted that early successional cover and edges would be used more
than expected during night and crepuscular periods associated with presumed
peaks in Bobcat foraging activities.
Field Site Description
The Joseph W. Jones Ecological Research Center at Ichauway was an
11,735-ha research facility located in Baker County, 16 km south of Newton,
GA. Approximately 24 km of the Ichawaynochaway Creek flowed through
the study area, and almost 22 km of the Flint River formed the eastern
boundary (Boring 2001). Elevations ranged from 27 to 61 m. The climate
was characterized by hot, humid summers and short, mild, wet winters, with
average daily temperatures ranging from 11 °C (winter) to 27 °C (summer).
Average annual precipitation was 132 cm per year (Boring 2001).
Pinus palustris P. Mill (Longleaf Pine) woodlands and limesink wetlands
were the dominant habitat types at Ichauway. Other habitats included mixed
pine-hardwood areas, food plots, agricultural fields, P. elliottii Enghelm
(Slash Pine) flatwoods, riparian hardwood hammocks, and Quercus spp.
(oak) sandhill barrens. Natural and old-field P. taeda L. (Loblolly Pine)
stands, Nyssa spp. (tupelo) trees, limesink ponds, creek swamps, forested
wetlands, riverine areas, shrub-scrub upland, and human/cultural (i.e.,
resident quarters) areas were also found at Ichauway (Boring 2001). The
understory was dominated by Aristida stricta Michx (Wiregrass) and oldfield grasses (e.g., Andropogon spp.), but >1000 vascular plant species
occurred on the site (Drew et al. 1998, Goebel et al. 1997).
Quail management practices were used at the study site, and included food
plots consisting of Brachiaria ramose Stapf (Brown Top Millet), Triticum
aestivum L. (Winter Wheat), Vigna spp. (cowpea), Sorghum vulgare Persoon
(Grain Sorghum), and Sorghum spp. (Egyptian wheat) (Godbois et al. 2004).
Supplemental feeding for quail with Grain Sorghum occurred on approximately
60% of the study area each year between November and May (Godbois
et al. 2004). Fields were disced to improve quail food availability, and limited
predator removal was practiced. Bobcats were not harvested during the study.
2010 J.D. Kirby, J.C. Rutledge, I.G. Jones, L.M. Conner, and R.J. Warren 319
Prescribed burning was performed on a 2-year rotation, usually during
winter and early spring, on approximately 4000 to 6000 ha throughout the
entire site (Godbois et al. 2004). Prescribed burning was used to control understory
vegetation, reduce hardwoods, manage wildlife habitat, reduce fuel
buildup, promote Wiregrass seed production, prepare sites for pine regeneration,
and for experimental research and educational activities (Boring 2001).
Bobcat capture, handling, and radiotelemetry
We trapped Bobcats using Victor No. 1.75 offset and No. 3 Soft Catch®
traps (Woodstream, Lititz, PA) from December 2000–May 2004. Captured
animals were netted and administered an intramuscular injection of ketamine
hydrochloride (10 mg/kg body weight; Seal and Kreeger 1987). We
classified animals as adult or juvenile (<1 year) based on secondary sex
characteristics, length, and weight (Crowe 1975). We fitted adults with a
180-g VHF radiocollar (Advanced Telemetry Systems, Isanti, MN); some
VHF radiocollars were outfitted with motion sensors. Each Bobcat received
a uniquely numbered ear tattoo. Bobcats were allowed to recover in portable
pet kennels placed in a secluded barn or other covered structure. They were
kept and monitored for 8–24 hr after immobilization to ensure full recovery
before release at the trap site. All trapping procedures were approved by
the University of Georgia Institutional Animal Care and Use Committee
Using radiotelemetry, we began monitoring Bobcats 2–7 d after release.
We obtained locations by triangulation, taking >2 radiotelemetry azimuths
from known reference points with a 3-element Yagi antenna (Sirtrack, New
Zealand) and hand-held receiver (Wildlife Materials, Carbondale, IL). We
located each Bobcat 4–6 times per wk, with >8 hr between each location.
To ensure equal sampling throughout the diel period, we shifted the start
time of each telemetry data collection period by 2 hours each wk. We followed
this diel schedule for each season. We performed accuracy tests for all
field personnel, and the standard deviation associated with bearing accuracy
was 7°. To minimize error due to animal movement between readings, we
attempted to get close to each animal and limited time between consecutive
bearings to <15 min, though most consecutive bearings were taken within 7
min (Cochran 1980, Kenward 1987, White and Garrott 1990).
We determined activity (active or inactive) by a change in the motion
switch of the transmitter or, on collars lacking the motion-sensitive switch,
by a change in signal intensity (i.e., fluctuation in signal pitch or strength)
when movement was detected (Chamberlain et al. 1998, Patterson et al.
1999). We defined an animal as active if the transmitter motion switch
was triggered or if the transmitter exhibited a change in signal intensity
while triangulating (Amstrup and Beecham 1976, Chamberlain et al. 1998,
Theuerkauf et al. 2003). Radiocollared Bobcats were distributed throughout
the study area.
320 Southeastern Naturalist Vol. 9, No. 2
We used the FORTRAN program EPOLY (L.M. Conner, Joseph W.
Jones Ecological Research Center, Newton, GA, unpubl. data) to convert
radiotelemetry bearings into Universal Transverse Mercator coordinates.
We calculated 95% minimum convex polygon (MCP; Mohr 1947) seasonal
(e.g., fall = 20 Sep–20 Dec; winter = 21 Dec–20 Mar; spring = 21 Mar–19
Jun; summer = 20 Jun–19 Sep) home ranges for Bobcats with ≥30 locations
per calendar season using CALHOME (Kie et al. 1996).
We performed cover-type analyses using ArcGis software (ESRI 2004). We
identified habitat using photo interpretation, classified habitat into eight cover
types (agriculture/food plot [20.2%], shrub/scrub [1.6%], hardwood [10.8%],
pine regeneration [4.4%], pine [31.8%], mixed pine-hardwood [24.5%],
wetland [5.0%], and urban/barren [1.5%]), and digitized habitat to create a
cover-type layer. We intersected Bobcat locations and home ranges onto the
cover-type layer using ArcGis (ESRI 2004).
We determined habitat use at Johnson's third order of selection (habitat
selection within an individual’s home range) to investigate whether activity
status and time of day affected habitat selection seasonally (Johnson 1980).
We considered daytime as 2 hr after sunrise until 2 hr before sunset, and
nighttime as 2 hr after sunset until 2 hr before sunrise. Finally, we defined
crepuscular periods as the time period 2 hr before and 2 hr after sunrise and
sunset. To simplify calculations, we assumed sunrise and sunset for each
day of a season occurred at the same time, determined as the time of sunrise
and sunset on the median day of each calendar season. We accounted for
Daylight Saving Time during the appropriate seasons.
We used a Euclidean distance technique to test for habitat selection by
comparing the mean distances between animal locations and cover types to
corresponding expected distances (Conner and Plowman 2001, Conner et
al. 2003). We generated random locations for each home range using the
random-point generator in the ArcGis Animal Movement Extension, and we
calculated the mean distance (m) from random locations within each Bobcat
home range to each cover type using the NEAR command in ArcGis (ESRI
2004). We also calculated mean distances from each Bobcat location to each
cover type in the home range. For third-order habitat selection, we created
eight distance ratios for each Bobcat (one for each cover type). Distance
ratios were calculated as the average distances from Bobcat locations within
the home range (i.e., “used” distances) to each cover type divided by the average
distances from random locations within the home range (i.e., random
distances) to each cover type.
We assessed third-order habitat selection as a function of sex, season,
time of day, and activity status using a four-factor MANOVA with year
treated as a block. We expected the ratio of used distances to random distances
to equal 1.0 if habitat use was random (i.e., no selection; Conner and
Plowman 2001, Conner et al. 2003). If the ratio was <1, then animals were
closer to the cover type than expected; whereas, if the ratio was >1, then the
animals were farther from the cover than expected. Thus, we considered
2010 J.D. Kirby, J.C. Rutledge, I.G. Jones, L.M. Conner, and R.J. Warren 321
cover types to be preferred if the ratio was <1 and avoided if the ratio was
>1. We used univariate t-tests to identify preferred (i.e., animal closer than
expected) and avoided (i.e., animal farther away than expected) cover types
if the MANOVA indicated selection was nonrandom. Finally, we ranked
cover types using ranking matrices (Conner and Plowman 2001, Conner et
al. 2003). All statistical analyses were performed with SAS software (SAS
Institute 2003). We considered statistical significance to occur at α ≤ 0.10.
The number of Bobcats monitored ranged from 13–27 individuals
seasonally from 21 September 2001 to 20 June 2004 for 11 consecutive
calendar seasons (43 total animals: 16 male [M] and 27 female [F]), and
we used 12,520 telemetry locations for analysis. Number of individual
locations ranged from 156–334 (SE = 7.38) per animal. For analysis of
cover-type selection and time of day, we used 7088 daytime telemetry
locations and 4332 nighttime telemetry locations. To analyze cover-type
selection and activity status, we used 6420 active animal locations and
5000 inactive animal locations. Bobcat cover-type selection did not differ
by sex (F8, 195 = 1.37, P = 0.2102), season (F80, 1245 = 1.08, P = 0.3056),
or their interaction (F80, 1245 = 0.60, P = 0.9937). Although there were no
significant two-way interactions (P > 0.10, all cases), habitat selection occurred
(Λ = 0.017, P = 0.001) when data for sex and season were pooled.
Bobcats were farther than expected from pine regeneration, pine, and
mixed pine-hardwood cover types, but were closer than expected to wetland
and urban/barren cover types (Table 1). Urban/barren cover was most
preferred, followed by wetland, hardwood, agriculture, shrub/scrub, pine
regeneration, pine, and mixed pine-hardwood (Table 2). We found no evidence
that activity status (Λ = 0.990, P = 0.981) or time of day (Λ = 0.972,
P = 0.647) affected Bobcat cover-type selection.
Table 1. Cover type distance ratios for third-order selection using seasonalA home ranges for
Bobcats at Ichauway, Baker County, GA, 2001–2004.
Cover type ρB t PC
Agriculture 0.037 0.75 0.456
Shrub/scrub 0.044 1.41 0.167
Hardwood 0.027 0.57 0.575
Pine regeneration 0.112 2.98 0.005
Pine 0.133 1.77 0.084
Mixed pine-hardwood 0.254 2.44 0.019
Wetland -0.000 -0.02 0.988
Urban/barren -0.043 -2.01 0.051
ASeasons were delineated according to calendar year (i.e., fall = 20 Sep–20 Dec, winter = 21
Dec–20 Mar, spring = 21 Mar–19 Jun, summer = 20 Jun–19 Sep).
BAverage distance from random locations within home ranges divided by average distance from
random locations throughout study area. Mean ratios <1 indicate cover type preference, and
>1 indicate habitat avoidance.
CProbability that the mean ratio = 1.0.
322 Southeastern Naturalist Vol. 9, No. 2
Bobcats selected cover types within their home ranges, but no sex-specific or seasonal differences were detected at this spatial scale. Bobcats used
agriculture, shrub/scrub, hardwood, and wetland cover types as expected,
but they were farther than expected (i.e., avoided) from pine regeneration,
pine, and mixed pine-hardwood cover types. However, Bobcats were closer
than expected to urban/barren cover, probably because several animals
maintained home ranges in close proximity to the laboratory, research, and
residential buildings on Ichauway, the location of most of the urban/barren
cover on site.
Bobcats likely selected cover types based on prey abundance and
availability. Sigmodon hispidus Say and Ord (Cotton Rat), a primary Bobcat
prey species in the Southeast, are typically most abundant in early to
mid-successional habitats dominated by grass/forb-shrub vegetation in
the understory (Boyle and Fendley 1987). Agricultural fields (i.e., food
plots and their associated edges) also attract and potentially concentrate
rodents such as Cotton Rats (Cummings and Vessey 1994). Hardwood
cover types contain a dense herbaceous understory and shrub interspersion
necessary to provide essential cover for prey species (Golley et al.
1965, Schnell 1968).
Supplemental grains were distributed throughout portions of the study
area for quail, which potentially concentrated Bobcat prey such as Cotton Rats
and may have influenced cover-type selection. Although a previous study at
Ichauway found that Bobcats were located closer to agricultural habitats in
which supplemental food for quail was distributed (Godbois et al. 2004), diet
studies suggested that quail comprised only 1.8% of Bobcat diet, whereas
rodents comprised 75.1% (Doughty 2004). Cover-type selection by Bobcats
on our study site were consistent with other studies in the Southeast, in which
Table 2. Rankings based on pair-wise comparisons between cover-type distance ratios for thirdorder
habitat selection, using seasonal home ranges of Bobcats monitored on Ichauway, Baker
County, GA, 2001–2004. Mixed = mixed pine-hardwood.
Urban/ Shrub/ Pine
barren Wetland Hardwood Agriculture scrub regeneration Pine Mixed
Urban/Barren + +++A + +++ +++ +++ +++
Wetland -B + B + + +++ +++ +++
Hardwood ---A - + + +++ + +++
Agriculture - - - + + + +++
Shrub/Scrub --- - - - + + +++
Pine --- --- --- - - + +
Pine --- --- - - - - +
Mixed --- --- --- --- --- - -
AThree plus signs indicate row cover type significantly preferred over column cover type and
three minus signs indicate column cover type significantly preferred over row cover type (ttest,
P < 0.10).
BA single plus or minus sign indicates that the row and column habitats have similar rankings.
2010 J.D. Kirby, J.C. Rutledge, I.G. Jones, L.M. Conner, and R.J. Warren 323
Bobcats preferred agricultural areas (Cochrane 2003, Conner et al. 1992) and
bottomland hardwoods (Cochrane 2003, Heller and Fendley 1986).
Cotton Rats, other rodents, and lagomorphs are typically most active during
crepuscular periods (Buie et al. 1979, Hall and Newsom 1976, Kitchings
and Story 1978, Marshall and Jenkins 1966). Bobcats have been documented
as most active during crepuscular periods, corresponding with activity peaks
of their primary prey species (Anderson 1987, Buie et al. 1979, Chamberlain
et al. 1998, Hall and Newsom 1976, Kitchings and Story 1978, Marshall
and Jenkins 1966). Because Bobcats tend to be more active when their prey
are most active, we expected active animals to select agriculture and other
early to mid-successional cover types more than inactive animals. We also
expected Bobcats to prefer early to mid-successional cover types during
crepuscular periods. Because our data were collected equally throughout the
diel period, we were able to assess the importance of these sources of variation
on the behavior of Bobcats.
Contrary to our prediction, we found no evidence that activity status or
time of day affected Bobcat cover-type selection, suggesting that cover types
used for foraging also contained suitable shelter and loafing areas. Therefore,
on our study area and perhaps elsewhere throughout Bobcat range, we
suggest that daytime-only radiomonitoring of our study animals would have
been adequate for studying cover-type selection. We speculate that Bobcats
did not have to move among cover types to meet refugia and foraging requirements.
An analysis of Bobcat time-specific and activity-specific habitat
selection on study areas with intrinsic differences in prey abundance among
cover types would provide a test of this hypothesis. Future studies to document
prey densities in various cover types on our study site would provide
evidence to support our suggestion that prey abundance was likely the main
factor influencing Bobcat cover-type selection. Finally, we recommend
multi-species research and perhaps reanalysis of existing radiotelemetry
data sets collected for other species to determine the pervasiveness of temporal
differences in cover-type selection.
Funding and other support were provided by the Joseph W. Jones Ecological
Research Center, University of Georgia, and the Georgia Department of Natural
Resources. M.J. Chamberlain, S.B. Castleberry, R.L. Hendrick, Jr., C. Nielsen,
and 2 anonymous reviewers provided valuable comments to earlier versions of
this manuscript. We thank the Jones Center Wildlife Lab, especially B. Rutledge,
M. Perkins, A. Subalusky, B. Howze, and A. Reid, as well as J. Wade, R. Varnum,
and B. Cross, for trapping assistance. Several Jones Center personnel, especially
B. Bass, M. Melvin, J. Brock and A. Sheffield, also provided critical help with
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