Prey Selection by Three Mesopredators that are Thought
to Prey on Eastern Wild Turkeys (Meleagris gallopavo
sylvestris) in the Pineywoods of East Texas
Haemish I.A.S. Melville, Warren C.Conway, Michael L.Morrison, Christopher E. Comer, and Jason B. Hardin
Southeastern Naturalist, Volume 14, Issue 3 (2015): 447–472
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22001155 SOUTHEASTERN NATURALIST 1V4o(3l.) :1444,7 N–4o7. 23
Prey Selection by Three Mesopredators that are Thought
to Prey on Eastern Wild Turkeys (Meleagris gallopavo
sylvestris) in the Pineywoods of East Texas
Haemish I.A.S. Melville1,2,*, Warren C.Conway3, Michael L.Morrison1,
Christopher E. Comer4, and Jason B. Hardin5
Abstract - Predation, especially during the nesting and poult-rearing seasons, may inhibit
Meleagris gallopavo (Wild Turkey) recruitment in east Texas. Numerous authors cite Lynx
rufus (Bobcat), Canis latrans (Coyote), and Procyon lotor (Raccoon) as predators of Wild
Turkey. Consequently, we investigated prey selection of these 3 common mesopredators
using scat analysis. We also investigated prey-population dynamics using capture–mark–
recapture techniques for small mammals (Rodentia), and spotlight surveys and track plate
counts for Sylvilagus floridanus (Eastern Cottontail). We found no evidence that mesopredators
preyed upon Wild Turkeys. Small mammals and lagomorphs were the primary
components of mesopredator diets. Small-mammal numbers varied seasonally; however,
Cottontail relative abundance did not. Mesopredator diets were most diverse in summer.
In summer, Bobcats increased their use of small mammals, whereas Coyotes and Raccoons
diversified their diets to include seasonal fruits. Decline in small-mammal populations and
increase in mesopredator dietary diversity coincided with Wild Turkey nesting and poultrearing
seasons, which potentially could result in an increased threat to Wild Turkeys during
the nesting and poult-rearing season.
Introduction
Meleagris gallopavo L. (Wild Turkey) used to occupy approximately 12 million
hectares in East Texas (Campo and Dickson 1990). Overharvesting and habitat
destruction led to their local extinction by 1883 (Campo and Dickson 1990, Newman
1945). Restoration attempts in this region have been ongoing since 1924
(Newman 1945). Although most attempts to reestablish populations of Wild Turkeys
in the United States have been successful, this has not been the case in east
Texas (Boyd and Oglesby 1975, Isabelle 2010, Lopez et al. 2000, Newman 1945)
where predation by mesopredators including Lynx rufus Schreber (Bobcat), Canis
latrans Say (Coyote), and Procyon lotor L. (Raccoon) are thought to confound reintroduction
programs of Wild Turkeys (George 1997, Kelly 2001).
Predation is considered the primary cause of mortality for Wild Turkeys (Hamilton
and Vangilder 1992, Hughes et al. 2005, Kennamer 2005, Miller and Leopold
1Department of Wildlife and Fisheries Sciences, Texas A&M University, College Station,
TX 77843. 2Current address - No. 8 25th Street, Zakher, Al Ain, United Arab Emirates. 3Department
of Natural Resources Management, Texas Tech University, MS 2125, Lubbock,
TX 79409-2125. 4Arthur Temple College of Forestry and Agriculture, Stephen F. Austin
State University, Nacogdoches, TX 75962. 5Texas Parks and Wildlife Department, Oakwood,
TX 75855. *Corresponding author - haemish.melville@gmail.com.
Manuscript Editor: Renn Tumlison
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1992). Female Meleagris gallopavo sylvestris Vieillot (Eastern Wild Turkeys), their
eggs, and poults are particularly susceptible to predation during nesting and brood
rearing (Lopez et al. 2000, Speake 1980). Predation may limit recruitment potential
of low-density populations (Kot et al. 1993, Messier and Crête 1985, Newsome et
al. 1989, Terborgh et al. 2001, Trout and Tittensor 1989), such as Wild Turkeys in
East Texas (Isabelle 2010).
Coyotes, Bobcats, and Raccoons live sympatrically over large portions of their
range, including east Texas. Bobcats are carnivorous (Anderson 1985, Anderson
and Lavallo 2003), whereas Coyotes and Raccoons are omnivorous (Schmidly
2004). Small mammals (Bartoszewicz et al. 2008, Fedriani et al. 2000, Litvaitis and
Harrison 1989) and lagomorphs (Azevedo et al. 2006, Bekoff and Gese 2003, Gehrt
2003, Schmidly 2004) are primary components of the diets of these mesopredators.
Behavioral mechanisms, including diet selection and use of space facilitate
co-existence among mesopredators (Wang and Macdonald 2009). Predators adapt
their feeding habits to the availability of suitable prey (Krebs 1978, Sunquist and
Sunquist 1989).
Several mechanisms affect mesopredator prey selection. Predators display functional
(Baker et al. 2001, Boutin 1995, Holling 1959) as well as behavioral and
numerical responses (Dunn 1977; Jędrzejewska and Jędrzejewski 1998; Schmidt
and Ostfeld 2003, 2008) to changes in prey availability. Temporal variation in
resource availability is a feature of the natural environment (Holt 2008). Smallmammal
populations vary seasonally (O’Connell 1989) and annually (Windberg
1998). It seems that mesopredators respond to variations in prey availability, consuming
preferred prey when they are available and diversifying their diets when
preferred prey are scarce (Schoener 1971) .
To gauge the effect of predation by mesopredators on Eastern Wild Turkeys in
the Pineywoods of east Texas, we investigated the following hypotheses: (1) Eastern
Wild Turkeys contribute significantly to the diets of Bobcats, Coyotes, and
Raccoons; (2) mesopredator diets vary seasonally; (3) mesopredator diets overlap
significantly; (4) mesopredator prey availability varies seasonally; (5) predators
respond functionally to changes in prey availability; and (6) low points of prey
populations coincide with the Wild Turkey nesting season. Our results will contribute
to the understanding of the threat posed by these mesopredators to Eastern Wild
Turkeys in east Texas.
Study area
We conducted our study on 2 sites from January 2009 to September 2011 in
the Pineywoods Ecoregion of Nacogdoches and Angelina counties, east Texas.
Study site 1 (31°30'N, 94°43'W) encompassed 1360 ha and was intensively
managed for wildlife using controlled burns on a 5-year rotational basis. Study
site 2 (31°25'N, 94°23'W) encompassed 5000 ha and was managed for timber,
with fire excluded from the system. We chose these study sites because they
were areas where stable Wild Turkey populations existed and they represented
major land uses in east Texas.
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H.I.A.S. Melville, W.C.Conway, M.L.Morrison, C.E. Comer, and J.B. Hardin
2015 Vol. 14, No. 3
The Pineywoods occur in the western Gulf coastal region and extend through
east Texas, northwestern Louisiana, and southwestern Arkansas. Much of the natural
vegetation of the Pineywoods has been altered due to planting of Pinus taeda
L. (Loblolly Pine) and the exclusion of fire (Omernik et al. 2008). Private land
holdings account for 63% of the Pineywoods, much of which is in small parcels
(0.4 to 3.6 ha). A consequence of this pattern of ownership is an increase in forest
fragmentation (US Department of Agriculture 2002). Pineywoods forests that are
not under silviculture typically consist of an overstory that includes pine and oak
species in association with Liquidambar styraciflua L. (Sweet Gum), Carya spp.
(hickories), Fraxinus americana L. (White Ash), and Ilex opaca Ait. (American
Holly) and an understory including Ilex vomitoria Ait. (Yaupon), Cornus florida L.
(Flowering Dogwood), Asimina triloba (L.) Dunal (Pawpaw), Callicarpa americana
L. (American Beautyberry), Chasmanthium laxum var. sessiliflorum (L.)
Yates (Longleaf Uniola), Rubus plicatus Weihe and Nees (Bramble Blackberry),
and grasses including Panicum spp.
Mean annual rainfall in the Pineywoods is 1192 mm, and monthly mean precipitation
varies from a minimum of 55 mm in July to a maximum of 116.4 mm in
May. The mean annual minimum and maximum temperatures are 12.8 °C and 25.5
°C, respectively, with a mean summer maximum of 35 °C (Sivanpillai et al. 2005).
Methods
We used the natural (delineated by solstices and equinoxes) seasons for all
analyses (winter: 21 December to 20 March, spring: 21 March to 20 June, summer:
21 June to 20 September, fall: 21 September to 20 December) because this schedule
is relevant to the habits of mesopredators and prey species, including Eastern Wild
Turkeys. We surveyed from winter 2009 (January) to summer 201 1 (August).
Mesopredator dietary analysis
To assess mesopredator dietary composition, we analyzed scat (Putman 1984)
collected from Coyotes, Bobcats, and Raccoons. Scats collected from these species
can be distinguished by appearance (Toweill and Anthony 1988). We searched
several drainage lines and creek beds once every 2 weeks, and collected scats
opportunistically on roads within study sites for the duration of the study. Scat collection
was non-random with respect to microhabitat (Neale and Sacks 2001).
After collection, we washed the scats and separated the contents into 4 categories:
(1) bones and teeth, (2) hair and feathers, (3) plant material, and (4) insects.
We identified scat-sample contents using osteological and hair keys, samples
from the study sites, and reference books (Toweill and Anthony 1988). In addition,
the microscopic characteristics of hairs were used to identify prey species
(Prugh 2005). We made cuticular-scale impressions on microscope slides (Melville
et al. 2004), examined them under compound microscopes, and compared
them to keys and reference slides to identify the taxa represented by the hair. Hair
medullary shapes from scat samples were compared to a key (Debelica and Thies
2009), and tooth and bone remains to reference material. When identificaton of
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samples could not be determined using standard references, we consulted specialist
taxonomists.
Prey population monitoring
Small mammals. Live-trapping grids (100 traps, 100 m x 100 m) and capture–
mark–recapture (CMR) methodologies were used to estimate small-mammal
populations (Edalgo and Anderson 2007, Parmenter et al. 2003, Reed et al. 2007,
Wiewel et al. 2007). Annually, we randomly selected 6 of a total of 29 one-yearold
Eastern Wild Turkey nest locations (3 on each study site) for grid placement.
Nests were located during a study investigating the nesting ecology of Eastern
Wild Turkeys in the Pineywoods of east Texas that ran concurrently with ours.
We also selected 6 random locations (3 on each study site) for a total of 12 CMR
surveys every season; 24 survey grids for the study. We sampled the same grids
seasonally and changed grid locations annually to ensure that the Wild Turkey
nest sites were a maximum of 1 year old. A mixture of rolled oats and peanut butter
was used as bait, and each grid was sampled for 7 consecutive nights. Each
animal that was captured was implanted with a uniquely numbered passive integrated
transponder (PIT) tag (Biomark®).
Our small-mammal capture records were used to estimate population size of the
3 most abundant small-mammal species: Reithrodontomys fulvescens J.A. Allen
(Fulvous Harvest Mouse), Sigmodon hispidus Say and Ord (Hispid Cotton Rat),
and Peromyscus leucopus Rafinesque (White-footed Mouse), for each trapping grid
using the robust design method in program MARK (White and Burnham 1999). To
ensure independence of samples, we positioned trapping grids in year 2 at least 1
km from any previous grid locations, and the grids on nest sites were associated
with new Wild Turkey nests.
Lagomorphs. We used spotlight counts and track-plate surveys to monitor
seasonal variation in abundance of Sylvilagus floridanus J.A. Allen (Eastern Cottontail)
(Malaney and Frey 2006, Williams et al. 2012). The 2 complimentary
monitoring protocols were implemented seasonally because track plates have not
been validated to monitor changes in relative abundance of lagomorphs (Ray and
Zielinski 2008).
A spotlight route was established on each study site. Routes traversed the length
and breadth of each study site. Each spotlight count was conducted by 2 observers,
each equipped with a spotlight and were initiated immediately after sunset,
counting all observable Cottontails. Each spotlight route was approximately 20 km
long (19 km on study site 2, 22 km on study site 1). We drove each route on alternate
evenings for 14 days (7 iterations per study site) each season and conducted
consecutive spotlight counts on individual study sites travelling in the opposite direction
to the previous count. Each route was driven at a constant speed of 10 km/h.
When bad weather curtailed counts, we delayed the spotlight surveys until the next
evening (Fletcher et al. 1999).
We conducted track-plate surveys comprising 25 track plates laid out in 2 parallel
lines of 12 and 13 on each site. We positioned track plates along the same
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2015 Vol. 14, No. 3
roads as those used during spotlight counts to facilitate direct comparison between
methods. Each track plate was located at least 150 m from the road. Track plates
were constructed by covering 1 side of plywood backing plates (0.5 m x 0.5 m)
with a Biofoam® impression surface (Hooper and Rea 2009). We placed track plates
at least 320 m apart (i.e., greater than 1 Eastern Cottontail home-range diameter;
Bond et al. 2002) so that each track plate could be considered an independent
sample (Hamm et al. 2003, Ray and Zielinski 2008). Track plates were deployed
simultaneously on both sites and exposed for 14 consecutive days. We checked the
track plates every alternate day (7 times during each exposure period) and recorded
any fresh track impressions. Fresh tracks, irrespective of whether they comprised
a single impression or several tracks, were recorded as a single detection for each
species on each track plate (Ray and Zielinski 2008, Sar geant et al. 1998).
Analyses
Mesopredator diet. We estimated relative frequency of each prey item within
mesopredator scats by species, study site, year, and season. We used chi-square
analysis (Zar 1999) to estimate differences among categories of dietary components
among and within species, between years, between sites, and among seasons
(Fedriani et al. 2000).
We calculated a Shannon-Wiener index of dietary diversity for each mesopredator
species (Jethva and Jhala 2004). Dietary overlap was calculated using Pianka’s
index (O; Fedriani et al. 2000, Glen and Dickman 2008, Pianka 1973) for each
species pair (Bobcat and Coyote, Bobcat and Raccoon, Coyote and Raccoon); we
considered overlap index values >0.6 to be biologically significant (Bethea et al.
2006, Pianka 1976, Wallace 1981).
Prey populations
Small mammals. We calculated minimum known alive (Krebs 1966, Merritt et
al. 2001) values for each species on each grid seasonally, and used factorial analysis
of variance (ANOVA; Zar 1999) to compare differences between years, seasons,
study sites, species, and site types (nest site versus random site). We utilized Pollock’s
Robust Design (Pollock 1982) in program MARK (White and Burnham
1999) to estimate populations of the 3 most commonly captured species: Fulvous
Harvest Mice, Hispid Cotton Rats, and White-footed Mice. Primary sampling intervals
were set at 7 consecutive days per season; secondary samples were individual
days within a primary period. We used these population estimates to compare species
by year, study site, season, and nest site versus random site using factorial
ANOVA (Zar 1999).
Lagomorphs. We used spotlight count data to calculate a spotlight count index,
and the number of Eastern Cottontail detections (R)/km to index relative abundance.
These index values were used to compare lagomorph abundance between
years, study sites, and seasons with factorial ANOVA (Zar 1999).
We calculated a track-plate index by dividing the number of track-plate detections
(RT) per unit time by number of track plates (TP) (Crooks 2002, Dijak and
Thompson 2000, Drennan et al. 1998, Glennon et al. 2002, Lenth et al. 2008,
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Wilson and Delahay 2001) on a site and seasonal basis. Index values between year,
season, and study site (Gentry and Vierling 2007) were compared using factorial
ANOVA (Zar 1999). We utilized a logistic regression model including the variables
year, study site, and season relative to detection of Eastern Cottontails on track
plates and validated it with Hosmer-Lemeshow and likelihood ratio tests. Linear
regression was used to determine if spotlight and track-plate indices correlated
(Schmidt et al. 2011).
We evaluated all statistical tests at α = 0.05.
Results
Mesopredator diets
We collected and analyzed 1764 mesopredator scats from January 2009 to
August 2011: Bobcat, n = 637 (fall = 27, spring = 160, summer = 152, winter =
298); Coyote, n = 841 (fall = 54, spring = 235, summer = 252, winter = 299); and
Raccoon, n = 286 (fall = 28, spring = 123, summer = 71, winter = 64). During this
study 3383 prey occurrences (Bobcat = 976, Coyote = 1688, Raccoon = 719) were
recorded (Melville 2012).
Bobcats. Forty types of ingesta were identified, comprising vegetable matter,
insects, fishes, birds, reptiles, and mammals (Appendix 1). Despite finding avian
remains in Bobcat scats, no Wild Turkey remains were found in these samples. Bobcat
diets did not differ between study sites (χ2 = 2.89, df = 21, P = 0.175). Bobcat
diets did not vary between 2010 and 2011 (χ2 = 16.21, df = 13, P = 0.238), but did
vary between 2009 and 2010 (χ2 = 22.92, df = 13, P = 0.043), effected largely by
differences in consumption of Odocoileus virginianus Zimmerman (White-tailed
Deer; 2009 = 10%, 2010 = 7.8%), lagomorphs (2009 = 24.3%, 2010 = 37.7%), and
small mammals (2009 = 49%, 2010 = 37.5%) including Hispid Cotton Rats (2009
= 28.4%, 2010 = 19.6%). Between 2009 and 2011, major differences in consumption
(χ2 = 47.98, df = 14, P < 0.001) were effected by insects (2009 = 1.4%, 2011
5.6%), White-tailed Deer (2009 = 10%, 2011 = 6.7%), lagomorphs (2009 = 24.3%,
2011 = 45.4%), and small mammals (2009 = 49%, 2011 = 27.7%) including Hispid
Cotton Rats (2009 = 28.4%, 2011 = 20.2%). Annual Bobcat diets did not differ from
fall or spring diets; however, they differed from summer and winter diets (Table 1).
Bobcat diets varied seasonally, with the greatest contribution of White-tailed Deer
(16.4%) in summer. Lagomorphs were the primary food source for Bobcats, with
the greatest contribution in the fall (41.1%). Hispid Cotton Rats were consistently
present in Bobcat diets, but contributed most (32.4%) in the winter. Some seasonal
fruits were detected in Bobcat diets—notably Vitis rotundifolia Michx. (Muscadine
Grape; 3%) in summer (Appendix 1). Spring and fall diets were similar (Table 1).
Diversity values for Bobcat diets varied (fall H = 1.93, summer H = 2.47; Table 2).
Coyotes. Forty-nine types of ingesta, including vegetable matter, insects, fishes,
birds, reptiles, and human-made goods (Appendix 2) were present in Coyote scats.
There were no Eastern Wild Turkey remains in Coyote scats, despite finding other
avian prey remains in samples. Coyote diets differed between sites (χ2 = 43.97,
df = 14, P < 0.001) and years (2009 vs 2010: χ2 = 75.37, df = 17, P < 0.001; 2009
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vs 2011: χ2 = 120.77, df = 17, P < 0.001; 2010 vs 2011: χ2 = 119.79, df = 17, P <
0.001). The dietary components that contributed most to the annual variation were
lagomorphs (2009 = 13.1%, 2010 = 12.4%, 2011 = 22.4%), and small mammals
(2009 = 19.5%, 2010 = 13.0%, 2011 = 7.7%), in particular Hispid Cotton Rats
(2009 = 11.8%; 2010 = 8.1%; 2011 = 5.1%). Annual diets differed from seasonal
diets, and seasonal diets differed (Table 1). Various dietary components peaked
Table 1. Chi-square test results comparing the seasonal diets of 3 mesocarnivores in the Pineywoods
of east Texas from January 2009 to August 2011.
Species Seasons compared χ2 df P
Bobcat Annual vs fall 19.44 14 0.149
Bobcat Annual vs spring 19.69 14 0.140
Bobcat Annual vs summer 39.51 14 <0.001
Bobcat Annual vs winter 29.30 14 0.009
Bobcat Fall vs spring 15.17 14 0.367
Bobcat Fall vs summer 27.18 12 0.007
Bobcat Fall vs winter 32.92 10 <0.001
Bobcat Spring vs summer 39.55 14 <0.001
Bobcat Spring vs winter 50.25 14 <0.001
Bobcat Summer vs winter 68.82 12 <0.001
Coyote Annual vs fall 68.79 19 <0.001
Coyote Annual vs spring 205.29 15 <0.001
Coyote Annual vs summer 165.66 19 <0.001
Coyote Annual vs winter 223.84 16 <0.001
Coyote Fall vs spring 211.73 15 <0.001
Coyote Fall vs summer 72.80 18 <0.001
Coyote Fall vs winter 152.12 13 <0.001
Coyote Spring vs summer 342.26 15 <0.001
Coyote Spring vs winter 260.34 13 <0.001
Coyote Summer vs winter 322.28 14 <0.001
Raccoon Annual vs fall 73.88 13 <0.001
Raccoon Annual vs spring 66.16 13 <0.001
Raccoon Annual vs summer 31.18 19 <0.001
Raccoon Annual vs winter 68.97 15 <0.001
Raccoon Fall vs spring 167.34 13 <0.001
Raccoon Fall vs summer 45.83 16 <0.001
Raccoon Fall vs winter 83.43 13 <0.001
Raccoon Spring vs summer 97.84 17 <0.001
Raccoon Spring vs winter 120.73 16 <0.001
Raccoon Summer vs winter 106.31 20 <0.001
Table 2. Shannon Wiener diversity index (H) values for the diets of 3 mesocarnivores in the Pineywoods
of east Texas from January 2009 to August 2011.
Season Bobcat (H) Coyote (H) Raccoon (H)
Annual 2.35 2.71 2.75
Fall 1.93 2.44 2.43
Spring 2.29 2.12 1.96
Summer 2.47 2.90 2.83
Winter 2.13 2.09 2.31
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seasonally in Coyote diets: White-tailed Deer (28.5%), Sus scrofa L.(Feral Hog;
18%), and Hispid Cotton Rats (12.6%) in winter, and lagomorphs (19.7%) in spring
(Appendix 2). Diversity values for Coyote diets varied seasonally (winter H = 2.08,
summer H = 2.90; Table 2).
Raccoons. Forty-one types of ingesta including vegetable matter, insects,
fishes, birds, mammals, reptiles, aquatic invertebrates, and human-made items
(Appendix 3) were present in Raccoon scats. No Eastern Wild Turkey remains were
found in Raccoon scats, despite there being remnants of other avian prey in the
samples. Raccoon diets did not differ between sites (χ2 = 19.03, df = 11, P = 0.061).
Raccoon diets differed between years (2009 vs 2010: χ2 = 23.17, df = 12, P = 0.026;
2009 vs 2011: χ2 = 45.34, df = 12, P < 0.001; 2010 vs 2011: χ2 = 67.62, df = 12, P <
0.001). Dietary components that contributed most to the annual variation included
insects (2009 = 14.3%, 2010 = 11%, 2011 = 35.3%), White-tailed Deer (2009 =
5.3%, 2010 = 12.7%, 2011 = 2.9%), lagomorphs (2009 = 7.3%, 2010 = 5%, 2011
= 15.4%), Hispid Cotton Rats (2009 = 7.3%, 2010 = 5.1%, 2011 = 2.9%), and
Cambarus sp (Crawfish) (2009 = 2%, 2010 = 2.5%, 2011 = 5.9%). Annual diets
differed from seasonal diets, and diets differed by season (Table 1). Various dietary
components peaked seasonally: insects in spring (24.5%) and winter (23.7%),
White-tailed Deer (12.7%) in winter, Hispid Cotton Rats in spring (7.5%) and summer
(7%), and Crawfish (6.8%) in winter. Diversity values for Raccoon diets varied
seasonally (spring H = 1.96, summer H = 2.83; Table 2).
Comparison of mesopredators dietary composition. Diets of Bobcats, Coyotes,
and Raccoons differed annually and seasonally (Table 3). Pianka’s overlap index values
indicated that diets of mesopredators overlapped to varying degrees of biological
significance (O > 0.6; Table 3). Bobcat diets overlapped greatly with Coyote diets annually
(O = 0.72) and seasonally (spring [O = 0.65] and summer [O = 0.68]). Bobcat
and Raccoon diets did not overlap. Coyote diets overlapped greatly with Raccoon
diets annually (O = 0.69) and seasonally (spring [O = 0.88] and summer [O = 0.76]).
Table 3. Chi-square test results and the associated Pianka dietary overlap (O) values for 3
mesocarnivores in the Pineywoods of east Texas from January 2009 to August 2011.
Species Compared Season χ2 df P O
Bobcat vs Coyote Annual 570.29 14 <0.001 0.72
Bobcat vs Coyote Fall 42.13 10 <0.001 0.58
Bobcat vs Coyote Spring 144.08 13 <0.001 0.65
Bobcat vs Coyote Summer 148.51 12 <0.001 0.68
Bobcat vs Coyote Winter 243.47 10 <0.001 0.58
Raccoon vs Coyote Annual 471.35 20 <0.001 0.69
Raccoon vs Coyote Fall 30.53 10 <0.001 0.30
Raccoon vs Coyote Spring 87.79 14 <0.001 0.88
Raccoon vs Coyote Summer 107.82 17 <0.001 0.76
Raccoon vs Coyote Winter 326.97 14 <0.001 0.40
Bobcat vs Raccoon Annual 746.07 20 <0.001 0.41
Bobcat vs Raccoon Fall 58.35 10 <0.001 0.19
Bobcat vs Raccoon Spring 206.39 14 <0.001 0.39
Bobcat vs Raccoon Summer 180.47 18 <0.001 0.45
Bobcat vs Raccoon Winter 344.24 11 <0.001 0.27
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Prey
Small mammals. We captured 1922 small mammals (representing 7 species) during
67,200 trap nights from January 2009 to December 2010 (Table 4). Number of
species captured varied between sample sites (F = 25.01; df = 5, 342; P < 0.001).
We captured 562 White-footed Mice, 42 Blarina brevicauda Say (Short-tailed
Shrew), 276 Hispid Cotton Rats, 863 Fulvous Harvest Mice, 76 Ochrotomys nuttalli
Harlan (Golden Mouse), 12 Neotoma floridana Ord (Eastern Woodrat) and 90
Peromyscus gossypinus LeConte (Cotton Mouse). We detected seasonal differences
in small-mammal captures (F = 16.37; df = 3, 344; P < 0.001; winter n = 855 vs
summer n = 330, Tukey HSD = -1.045, P < 0.001; winter n = 855 vs fall n = 250,
Tukey HSD = -1.393, P < 0.001; spring n = 678 vs summer n= 330, Tukey HSD =
-0.701, P = 0.009; spring n = 678 vs fall n = 250, Tukey HSD = -1.05, P < 0.001).
(Fig. 1). There was no variation in small-mammal captures between years (F =
0.117; df = 1, 346; P = 0.733), study sites (F = 0.108; df = 1, 346; P = 0.743), or
grid types (nest or random) (F = 2.233; df = 1, 346; P = 0.136).
Table 4. Number of small mammals captured during a capture–mark–recapture survey in the Pineywoods
of east Texas, from January 2009 to December 2010.
Common name Scientific name Total captures
White-footed Mouse Peromyscus leucopus 563
Short-tailed Shrew Blarina carolinensis 42
Hispid Cotton Rat Sigmodon hispidus 276
Fulvous Harvest Mouse Reithrodontomys fulvescens 863
Golden Mouse Ochrotomys nuttalli 76
Eastern Woodrat Neotoma floridana 12
Cotton Mouse Peromyscus gossypinus 90
Total 1922
Figure 1. Trends in seasonal numbers of small mammals captured during a mark–recapture
survey in the Pineywoods of east Texas, January 2009–December 2010.
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Population estimates for the 3 most commonly captured species of small mammals
did not vary between years (Fig. 2). Harvest Mouse populations were greatest
during winter (mean = 20.5, SE = 2.86) and spring (mean = 19.1, SE = 2.91), then
declined during the summer months (mean = 5.4, SE = 0.95) into fall (mean = 4.3,
SE = 0.97) (Table 5, Fig. 2). Numbers of White-footed Mice varied between study
sites (site 1 mean = 5.8, SE = 0.84 vs. study site 2 mean = 9.8, SE = 1.29). Whitefooted
Mice followed similar seasonal patterns to Harvest Mice, (winter mean =
14.3, SE = 1.83; spring mean = 7.76, SE = 1.37; summer mean = 4.0, SE = 0.73; and
fall mean =4.9, SE = 1.15; Table 5; Fig. 2). Hispid Cotton Rat populations varied by
grid type, with greater numbers occurring on random sites (mean = 7.6, SE = 1.36)
than on Wild Turkey nest sites (mean = 3.2, SE = 0.47). Numbers of Hispid Cotton
Rats varied seasonally (summer mean = 9.7, SE = 2.3; fall mean = 2.6, SE = 0.8;
winter mean = 3.4, SE = 2.3; and spring mean = 4.4, SE = 1.0; Table 5, Fig. 2).
Lagomorphs. We conducted 70 spotlight counts over 1433 km (study site 2: 769
km, study site 1: 664 km) seasonally from April 2010 to August 2011, and recorded
132 Eastern Cottontails (study site 2 = 100, study site 1 = 32; Table 6). We observed
no difference in numbers of Eastern Cottontails between years (F = 0.025; df = 1,
68; P = 0.874) or seasons (F = 1.56; df = 3, 66; P = 0.207). Spotlight indexes differed
between sites (study site 2: mean = 0.2 R/km, SE = 0.02; study site 1: mean
= 0.04 R/km, SE = 0.01; F = 23.27; df = 1, 68; P < 0.001; Fig. 3).
Figure 2. Mean population numbers (± 1 SE) for the 3 most abundant small mammals captured
in the Pineywoods of east Texas, January 2009–December 2010.
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We recorded 3500 track-plate nights for lagomorphs from April 2010 to August
2011, and detected Eastern Cottontails 121 times (study site 2: n = 73; study
site 1: n = 48). We converted these to an index of detections per track plate (RT/
TP, range = 0.0–0.2; Table 7). There was no difference in track-plate indexes between
years (F = 0.87; df = 1, 68; P = 0.354) or seasons (F = 1.63; df = 3, 66; P =
0.191), but we detected a difference between sites (study site 2: mean = 0.08 RT/
TP, SE = 0.01; study site 1: mean = 0.06 RT/TP, SE = 0.01; F = 4.371; df = 1, 68;
P = 0.04; Fig. 4).
We used logistic regression to model the influence of the variables year, study
site, and season on detection of lagomorph on the track-plates. Detection of tracks
was influenced by study site (Z = -2.344, df = 1, P = 0.091; Table 8). This finding
Table 5. Results of ANOVA comparing the variables associated with the 3 most abundant species
caught in the Pineywoods of east Texas from January 2009 to August 2011.
Harvest Mouse Hispid Cotton Rat White-footed Mouse
Comparisons F df P F df P F df P
Study sites 0.757 1, 77 0.39 1.081 1, 41 0.31 6.201 1, 81 0.02
Years 0.436 1, 77 0.51 1.105 1, 41 0.30 0.308 1, 81 0.50
Grid types 0.351 1, 77 0.56 5.914 1, 41 0.02 0.107 1, 81 0.74
Seasons 18.340 3, 75 <0.001 4.476 3, 39 0.01 12.190 3, 79 <0.001
Tukey P Tukey P Tukey P
Winter vs spring -0.274 0.92 -0.550 0.56 -0.953 0.02
Winter vs summer -2.241 <0.001 0.416 0.76 -1.733 <0.001
Winter vs fall -2.574 <0.001 -1.065 0.08 -1.599 <0.001
Spring vs summer -1.966 <0.001 0.968 0.12 -0.779 0.08
Spring vs fall -2.299 <0.001 -0.513 0.64 -0.646 0.20
Summer vs fall -0.333 0.87 -1.481 0.01 0.133 0.98
Table 6. Spotlight index values (rabbits per km) for detections of Sylvilagus floridanus (Eastern Cottontail
Rabbit) in the Pineywoods of east Texas from spring 2010 to summer 2011.
Site Season n SE Spotlight index
Both All 70 0.01 0.10
Site 1 All 35 0.02 0.15
Site 2 All 35 0.01 0.04
Both Fall 14 0.02 0.05
Site 1 Fall 7 0.03 0.04
Site 2 Fall 7 0.03 0.06
Both Spring 28 0.02 0.10
Site 1 Spring 14 0.02 0.06
Site 2 Spring 14 0.03 0.15
Both Summer 14 0.05 0.13
Site 1 Summer 7 0.01 0.01
Site 2 Summer 7 0.06 0.26
Both Winter 14 0.02 0.87
Site 1 Winter 7 0.02 0.04
Site 2 Winter 7 0.04 0.14
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was confirmed by a likelihood ratio test (log likelihood = -434.05, df = 5, P =
0.037), and a Hosmer-Lemeshow goodness-of-fit test (χ2 = 6.89, df = 8, P = 0.55).
There was no correlation between our spotlight and track-plate indices (F =
1.349; df = 1, 68; P = 0.25; r2 = 0.02).
Discussion
Bobcats, Coyotes, and Raccoons had diverse diets in the Pineywoods of east
Texas. Their diets included a variety of mammals, insects, birds, plants, fishes, and
reptiles, but no evidence of Eastern Wild Turkey remains. Our results support previous
studies in the Southeast that have found limited support for the supposition
Figure 3. Spotlight count index (rabbits/km) for 2 study sites in the Pineywoods of east
Texas, April 2010–August 2011.
Table 7. Track plate index (rabbit impressions per track plate) for detections of Sylvilagus floridanus
(Eastern Cottontail Rabbit) in the Pineywoods of east Texas from spring 2010 to summer 2011.
Site Season n SE Track index
Both All 70 0.01 0.07
Site 1 All 35 0.01 0.08
Site 2 All 35 0.01 0.06
Both Fall 14 0.01 0.05
Site 1 Fall 7 0.01 0.05
Site 2 Fall 7 0.02 0.06
Both Spring 28 0.01 0.06
Site 1 Spring 14 0.02 0.05
Site 2 Spring 14 0.02 0.07
Both Summer 14 0.02 0.08
Site 1 Summer 7 0.02 0.04
Site 2 Summer 7 0.02 0.13
Both Winter 14 0.01 0.09
Site 1 Winter 7 0.02 0.08
Site 2 Winter 7 0.01 0.10
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2015 Vol. 14, No. 3
that mesopredators prey on Wild Turkeys, even in areas with abundant populations
of this potential prey (Chamberlain and Leopold 1999, Leopold and Chamberlain
2002, Wagner and Hill 1994).
When mesopredators prey on large eggs, they seldom ingest entire egg shells
(Larivière 1999). Even if mesopredators consumed large numbers of eggs, it would
be unlikely that we would have found many eggshell fragments in scats. It is also
unlikely that we would have detected poults in scats (Wagner and Hill 1994). Thus,
despite the lack of evidence, it is possible that mesopredators preyed on Eastern
Wild Turkey eggs and poults. Numerous investigations of Wild Turkey mortality
indicate that predation is the most important cause of reduced survival (Isabelle
2010, Kurzejeski et al. 1987, Miller et al. 1998, Speake et al. 1985, Swank et al.
1985), yet there is conflicting evidence relating to the importance of predation by
mesopredators on Wild Turkeys. Our results in east Texas confirm the findings of
other studies that Wild Turkeys do not contribute greatly to the diets of mesopredators
(Chamberlain and Leopold 1999, Leopold and Chamberlain 2002, Melville
2012, Wagner and Hill 1994).
Figure 4. Eastern Cottontail Rabbit track index (tracks/plate/night) calculated for 2 study
sites in the Pineywoods of east Texas, April 2010–August 2011.
Table 8. Confidence intervals from logistic regression of the variables associated with likelihood of
detecting Sylvilagus floridanus (Eastern Cottontail Rabbit) tracks on track plates in the Pineywoods
of east Texas from Spring 2010 to Summer 2011. *** = highly significant; * = significant.
Variable Lower Upper Z df P
(Intercept) -2.838 -1.353 -5.54 1 <0.001 ***
Site = Site 1 -0.834 -0.077 -2.34 1 0.019 *
2011 -0.629 0.629 0.00 1 1.000
Spring -1.042 0.109 -1.56 1 0.118
Summer -0.929 0.711 -0.26 1 0.795
Fall -1.496 0.241 -1.41 1 0.159
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Small mammals contributed greatly to the diets of mesopredators in the Pineywoods
of east Texas, although their populations declined from spring into summer
during 2009 and 2010. During spring and summer, mesopredators increased diversity
in their diets. Increased dietary diversity has been linked to limitation in food
availability (Clavero et al. 2003). Decline in prey availability occurred while Wild
Turkeys were nesting and raising poults. The synchrony between the decline in
prey populations and nesting of Wild Turkeys may have increased the probability
of Wild Turkey nest and poult predation, especially considering that mesopredators
diversified their diets during this period. Despite this, we found no Wild Turkey
remains in mesopredator scats.
Sympatric predators display variability in dietary breadth and overlap consistent
with principles of resource partitioning (Azevedo et al. 2006). Our results for Bobcats,
Coyotes, and Raccoons in the Pineywoods confirms previous findings that the
degree of dietary overlap between mesopredators varies seasonally (Azevedo et al.
2006, Chamberlain and Leopold 1999, Fedriani et al. 2000, Major and Sherburne
1987). Levels of dietary overlap between Bobcats and Coyotes were marginally
less than what is considered biologically significant (O > 0.6) in fall and winter,
but overlap increased to biologically significant levels in spring and summer. Similar
patterns of seasonal variation in dietary overlap between Bobcats and Coyotes
have been observed in central Mississippi (Chamberlain and Leopold 1999) and
California (Fedriani et al. 2000). An even more pronounced seasonal change in
significance of overlap was evident between Coyotes and Raccoons. There was no
biologically significant dietary overlap in winter or fall; however, dietary overlap
was significant in spring and summer. This change in overlap was probably due
to their selection of seasonal fruits. Few other studies have investigated dietary
overlap between Coyotes and Raccoons. Where comparisons have been made, they
were conducted on an annual basis and provided results that showed little overlap
between Coyote and Raccoon diets (Azevedo et al. 2006). Although a seasonal
trend of increasing dietary overlap was evident between Bobcats and Raccoons, the
level of overlap was never biologically significant. The increase in dietary overlap
between mesopredators in spring and summer, combined with high dietary diversity
index values in summer may indicate seasonal resource limitation in terms of food
availability during those times of year (Clavero et al. 2003).
Recent weather events can affect small-mammal communities (French et al.
1976, Grant et al. 1985, Schmidt and Ostfeld 2008). Our study of small mammals
spanned 2009 and 2010, a period associated with the onset of a 500-year drought in
Texas from 2010–2011 (CIESS 2012). The mechanism of influence of harsh weather
on small-mammal populations occurs indirectly through the impact on vegetation
productivity (Grant et al. 1985). Despite this, we found no evidence of a general
decline in small-mammal populations in 2010. Our results are similar to those of
Grant et al. (1985) that found no effect of temperature and precipitation on Hispid
Cotton Rat and White-footed Mouse populations. Grant et al. (1985) also found that
Fulvous Harvest Mouse populations were influenced by temperature fluctuations,
but not precipitation. In the Pineywoods, Fulvous Harvest Mouse numbers declined
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2015 Vol. 14, No. 3
consistently during the warmer months. It is unlikely that the full effect on small
mammals of the Texas drought of 2010–2011 was fully identified during our study
because this would only have been evident after our monitoring ceased.
Bobcats, Coyotes, and Raccoons preyed on small mammals to varying degrees.
The only small-mammal species that contributed substantially to mesopredator
diets was the Hispid Cotton Rat, a primary prey item for Bobcats. Other small
mammals were preyed upon opportunistically by mesopredators. Mesopredators
are known to limit Hispid Cotton Rat populations, especially in areas where fire is
excluded (Conner et al. 2011). This may be the case in the Pineywoods, especially
on commercial timber sites where fire has been excluded as a management tool. It
is unknown whether mesopredators limit other small mammals.
Mesopredators seemed to respond to variation in prey availability. Bobcats
responded to changes in Hispid Cotton Rat availability by increasing their consumption
of them when the population increased, similar to what was observed in Georgia
(Baker et al. 2001). Lagomorphs were present in the diets of the mesopredators under
study. Coyotes and Bobcats used lagomorphs extensively, despite their apparent
low densities. Cottontails seemed to be the most important dietary item for Bobcats,
consistently comprising more than 20% of the items in their scats, although their frequency
seemed to vary relative to availability of Hispid Cotton Rats.
Coyotes and Raccoons are opportunistic, generalist predators, varying their
food intake relative to availability (Bekoff and Gese 2003, Gehrt 2003). During
spring and summer, Coyote and Raccoon diets contained a large percentage of
fruit. Locating and consuming fruit probably required less energy than searching
for and capturing live prey. Thus, Coyotes and Raccoons likely improved foraging
efficiency by including a high proportion of fruit when seasonally abundant (Chamberlain
and Leopold 1999, MacArthur and Pianka 1966, Norber g 1977).
Eastern Wild Turkeys did not contribute to diets of Bobcats, Coyotes, or Raccoons
in east Texas during our study. Our study did not prove that the predators
under study did not use Eastern Wild Turkeys, but our data did not reveal use
during our study. It seems that mesopredators responded to seasonal variations in
resource availability, and that Eastern Cottontails, despite occurring in seemingly
low densities, contributed greatly to their diets. We suggest that further studies be
undertaken to determine if the mesopredators in question prey extensively on nests
and poults of Eastern Wild Turkeys in the Pineywoods. Further, we suggest that
more investigations into the population dynamics of Eastern Cottontails in east
Texas are warranted.
Acknowledgments
Our project was funded by federal excise taxes on sport-hunting arms and ammunition,
Grant W132R, with the Texas Parks and Wildlife Department, as well as by Texas
A&M University (Department of Wildlife and Fisheries Science), Stephen F. Austin State
University (The Arthur Temple College of Forestry and Agriculture), and The Rumsey
Research and Development Fund. We thank all the researchers and research technicians
including A. Wadyko, J. van Woert, T. Yurick, J. Deatherage, J.Rogers, J. Isabelle, A. Davis,
S. Seidel, and J. Wisnant.
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2015 Vol. 14, No. 3
462
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Appendix 1. The occurrence of dietary items in the Lynx rufus (Bobcat) scats collected in the Pineywoods of east Texas, January 2009–August 2011.
* indicates the presence of Bobcat hair in their own scats, which was attributed to grooming.
Annual 2009 2010 2011 Fall Spring Summer Winter
Type/species Common name n % n % n % n % n % n % n % n %
Bird
Cardinalis cardinalis Cardinal 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2
Gallus gallus domesticus Chicken 24 2.5 16 2.5 6 2.9 2 1.7 4 10.3 5 2.0 7 3.0 8 1.8
Picoides sp. Woodpecker 1 0.1 0 0.0 1 0.5 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2
Unidentified bird 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 1 0.2
Fish
Unidentified fish 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 2 0.8 0 0.0 0 0.0
Insects
Unidentified Insects 19 2.0 9 1.4 3 1.5 7 5.9 2 5.1 12 4.9 1 0.4 4 0.9
Mammals
Odocoileus virginianus White tailed deer 88 9.1 64 10.0 16 7.8 8 6.7 3 7.7 26 10.6 38 16.4 21 4.7
Ovis aries Sheep 1 0.1 0 0.0 0 0.0 1 0.8 0 0.0 1 0.4 0 0.0 0 0.0
Sus scrofa Hog 12 1.2 8 1.3 3 1.5 1 0.8 0 0.0 4 1.6 2 0.9 6 1.3
Dasypus novemcinctus Armadillo 4 0.4 2 0.3 2 1.0 0 0.0 0 0.0 1 0.4 3 1.3 0 0.0
Lynx rufus Bobcat 9* 6* 2* 1* 0* 2* 2* 5*
Felis sylvestris catus Domestic Cat 3 0.3 2 0.3 1 0.5 0 0.0 0 0.0 1 0.4 0 0.0 2 0.4
Urocyon cinereoargenteus Gray Fox 7 0.7 4 0.6 1 0.5 2 1.7 0 0.0 2 0.8 1 0.4 4 0.9
Didelphis virginiana Opossum 3 0.3 2 0.3 1 0.5 0 0.0 0 0.0 1 0.4 0 0.0 2 0.4
Procyon lotor Raccoon 29 3.0 19 3.0 5 2.5 5 4.2 0 0.0 9 3.7 11 4.7 9 2.0
Vulpes vulpes Red Fox 2 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 1 0.4 0 0.0
Sylvilagus floridanus Eastern Cottontail 245 25.3 136 21.3 58 28.4 51 42.9 15 38.5 73 29.8 42 18.1 115 25.5
Sylvilagus aquaticus Swamp Rabbit 41 4.2 19 3.0 19 9.3 3 2.5 1 2.6 13 5.3 5 2.2 22 4.9
Sciurus niger Eastern Fox Squirrel 3 0.3 2 0.3 1 0.5 0 0.0 0 0.0 0 0.0 1 0.4 2 0.4
Sciurus carolinensis Eastern Gray Squirrel 1 0.1 0 0.0 1 0.5 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2
Neotoma floridana Eastern Woodrat 69 7.1 48 7.5 17 8.3 4 3.4 3 7.7 14 5.7 15 6.5 37 8.2
Sigmodon hispidus Hispid Cotton Rat 246 25.4 182 28.4 40 19.6 24 20.2 6 15.4 47 19.2 47 20.3 146 32.4
Rattus norvegicus Norway Rat 6 0.6 2 0.3 3 1.5 1 0.8 0 0.0 2 0.8 1 0.4 3 0.7
Rattus rattus Rat 2 0.2 0 0.0 0 0.0 0 0.0 1 2.6 0 0.0 0 0.0 1 0.2
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Annual 2009 2010 2011 Fall Spring Summer Winter
Type/species Common name n % n % n % n % n % n % n % n %
Peromyscus gossypinus Cotton Mouse 20 2.1 16 2.5 4 2.0 0 0.0 0 0.0 4 1.6 7 3.0 9 2.0
Peromyscus leucopus White-footed Mouse 21 2.2 15 2.3 3 1.5 3 2.5 1 2.6 6 2.4 2 0.9 12 2.7
Ochrotomys nuttalli Golden Mouse 7 0.7 6 0.9 1 0.5 0 0.0 0 0.0 0 0.0 2 0.9 5 1.1
Reithrodontomys humulis Eastern Harvest Mouse 20 2.1 19 3.0 1 0.5 0 0.0 0 0.0 3 1.2 9 3.9 8 1.8
Reithrodontomys fulvescens Fulvous Harvest Mouse 34 3.5 26 4.1 7 3.4 1 0.8 1 2.6 4 1.6 10 4.3 19 4.2
Snake
Coluber spp. Racer species 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0
Nerodia spp. Watersnake species 1 0.1 1 0.2 0 0.0 0 0.0 1 2.6 0 0.0 0 0.0 0 0.0
Agkistrodon piscivorus Cotton Mouth 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0
Crotalus horridus Timber Rattlesnake 1 0.1 0 0.0 1 0.5 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0
Plant
Callicarpa americana Beauty Berry 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0
Rubus plicatus Bramble Blackberry 2 0.2 1 0.2 0 0.0 1 0.8 0 0.0 2 0.8 0 0.0 0 0.0
Prunus virginiana Choke Cherry 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 0 0.0 2 0.9 0 0.0
Unidentified grass 35 3.6 21 3.3 9 4.4 5 4.2 1 2.6 11 4.5 11 4.7 12 2.7
Smilax rotundifolia Greenbriar 2 0.2 2 0.3 0 0.0 0 0.0 0 0.0 0 0.0 2 0.9 0 0.0
Vitis rotundifolia Muscadine 7 0.7 7 1.1 0 0.0 0 0.0 0 0.0 0 0.0 7 3.0 0 0.0
Pyrus spp. Pear 1 0.1 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0
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Appendix 2. The occurrence of dietary items in the Canis latrans (Coyote) scats collected in the Pineywood of east Texas, January 2009–August 2011.
* indicates the presence of Coyote hair in their own scats, which was attributed to grooming.
Annual 2009 2010 2011 Fall Spring Summer Winter
Type/species Common name n % n % n % n % n % n % n % n %
Bird
Cardinalis cardinalis Cardinal 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2
Gallus gallus domesticus Chicken 41 2.5 15 1.5 19 4.8 7 2.6 5 5.2 3 0.6 12 2.1 21 4.4
Zenaida macroura Mourning Dove 2 0.1 1 0.1 0 0.0 1 0.4 0 0.0 0 0.0 0 0.0 2 0.4
Unidentified bird 10 0.6 5 0.5 4 1.0 1 0.4 2 2.1 4 0.8 4 0.7 0 0.0
Fish
Unidentified fish 2 0.1 2 0.2 0 0.0 0 0.0 0 0.0 1 0.2 1 0.2 0 0.0
Insect
Unidentified Insect 70 4.2 25 2.5 10 2.5 35 12.9 0 0.0 55 10.6 11 2.0 4 0.8
Mammal
Odocoileus virginianus White-tailed Deer 292 17.7 170 17.2 70 17.8 52 19.1 22 22.7 79 15.3 55 9.8 136 28.5
Equus caballus Horse 1 0.1 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0 0 0.0 0 0.0
Sus scrofa Hog 145 8.8 92 9.3 26 6.6 27 9.9 6 6.2 26 5.0 27 4.8 86 18.0
Ovis aries Sheep 1 0.1 0 0.0 0 0.0 1 0.4 0 0.0 1 0.2 0 0.0 0 0.0
Lynx rufus Bobcat 5 0.3 3 0.3 2 0.5 0 0.0 0 0.0 1 0.2 4 0.7 0 0.0
Canis latrans Coyote 34* 21* 9* 4* 1* 13* 9* 11*
Urocyon cinereoargenteus Gray Fox 9 0.5 5 0.5 3 0.8 1 0.4 0 0.0 1 0.2 4 0.7 4 0.8
Didelphis virginiana Opossum 2 0.1 1 0.1 0 0.0 1 0.4 0 0.0 0 0.0 0 0.0 1 0.2
Procyon lotor Raccoon 38 2.3 16 1.6 12 3.1 10 3.7 1 1.0 9 1.7 15 2.7 13 2.7
Dasypus novemcinctus Armadillo 21 1.3 7 0.7 12 3.1 2 0.7 0 0.0 7 1.4 13 2.3 1 0.2
Lepus californicus Black-tailed Jack Rabbit 2 0.1 1 0.1 1 0.3 0 0.0 0 0.0 1 0.2 0 0.0 1 0.2
Sylvilagus floridanus Eastern Cottontail 218 13.2 120 12.2 43 10.9 55 20.2 11 11.3 94 18.2 43 7.7 70 14.7
Sylvilagus aquaticus Swamp Rabbit 21 1.3 9 0.9 6 1.5 6 2.2 0 0.0 8 1.5 7 1.2 6 1.3
Neotoma floridana Eastern Woodrat 33 2.0 21 2.1 9 2.3 3 1.1 1 1.0 10 1.9 8 1.4 14 2.9
Rattus norvegicus Norway Rat 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2
Sigmodon hispidus Hispid Cotton Rat 162 9.8 116 11.8 32 8.1 14 5.1 11 11.3 42 8.1 49 8.7 60 12.6
Peromyscus gossypinus Cotton Mouse 11 0.7 9 0.9 1 0.3 1 0.4 1 1.0 2 0.4 5 0.9 3 0.6
Peromyscus leucopus White-footed Mouse 9 0.5 6 0.6 3 0.8 0 0.0 0 0.0 2 0.4 3 0.5 4 0.8
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Annual 2009 2010 2011 Fall Spring Summer Winter
Type/species Common name n % n % n % n % n % n % n % n %
Ochrotomys nuttalli Golden Mouse 5 0.3 3 0.3 2 0.5 0 0.0 0 0.0 0 0.0 4 0.7 1 0.2
Reithrodontomys humulis Eastern Harvest Mouse 14 0.8 13 1.3 0 0.0 1 0.4 0 0.0 2 0.4 8 1.4 4 0.8
Reithrodontomys fulvescens Fulvous Harvest Mouse 27 1.6 21 2.1 4 1.0 2 0.7 3 3.1 3 0.6 10 1.8 11 2.3
Mus muscullus House Mouse 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0
Geomys breviceps Baird’s Pocket Gopher 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0
Plant
Callicarpa americana Beauty Berry 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0
Rubus plicatus Bramble Blackberry 174 10.6 114 11.6 21 5.3 39 14.3 0 0.0 141 27.3 33 5.9 0 0.0
Prunus virginiana Choke Cherry 37 2.2 26 2.6 11 2.8 0 0.0 3 3.1 0 0.0 34 6.1 0 0.0
Zea mays Corn 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0
Unidentified grasses 90 5.5 63 6.4 19 4.8 8 2.9 3 3.1 20 3.9 42 7.5 25 5.2
Smilax rotundifolia Green briar 58 3.5 40 4.1 17 4.3 1 0.4 7 7.2 1 0.2 49 8.7 1 0.2
Morus nigra Mulberry 9 0.5 5 0.5 4 1.0 0 0.0 6 6.2 0 0.0 3 0.5 0 0.0
Vitis rotundifolia Muscadine 86 5.2 54 5.5 31 7.9 1 0.4 5 5.2 0 0.0 81 14.4 5 1.0
Pyrus spp. Pear 1 0.1 0 0.0 1 0.3 0 0.0 1 1.0 0 0.0 0 0.0 0 0.0
Ampelopsis arborea Peppervine 5 0.3 0 0.0 5 1.3 0 0.0 0 0.0 0 0.0 5 0.9 0 0.0
Diospyros virginiana Persimone 21 1.3 11 1.1 10 2.5 0 0.0 8 8.2 0 0.0 13 2.3 0 0.0
Unidentified plants 8 0.5 0 0.0 8 2.0 0 0.0 0 0.0 0 0.0 8 1.4 0 0.0
Ligustrum vulgare Privet 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0
Cucurbita spp. Pumpkin 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 1 0.2 0 0.0
Citrullus lanatus Watermelon 4 0.2 1 0.1 2 0.5 1 0.4 0 0.0 0 0.0 4 0.7 0 0.0
Reptile
Unidentified lizard 1 0.1 0 0.0 1 0.3 0 0.0 1 1.0 0 0.0 0 0.0 0 0.0
Unidentified snakes 2 0.1 1 0.1 0 0.0 1 0.4 0 0.0 2 0.4 0 0.0 0 0.0
Miscellaneous
Leather 2 0.1 2 0.2 0 0.0 0 0.0 0 0.0 2 0.4 0 0.0 0 0.0
Latex 1 0.1 0 0.0 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2
Plastic 1 0.1 1 0.1 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.2
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Appendix 3. The occurrence of dietary items in the Procyon lotor (Raccoon) scats collected in the Pineywood of east Texas, January 2009–August 2011.
* indicates the presence of Raccoon hair in their own scats, which was attributed to grooming.
Annual 2009 2010 2011 Fall Spring Summer Winter
Type/species Common name n % n % n % n % n % n % n % n %
Bird
Gallus gallus domesticus Chicken 8 1.4 5 1.7 2 1.7 1 0.7 1 1.8 1 0.4 5 3.5 1 0.8
Picoides sp. Woodpecker 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8
Unidentified bird 5 0.9 2 0.7 1 0.8 2 1.5 0 0.0 2 0.8 1 0.7 2 1.7
Fish
Unidentified fish 4 0.7 3 1.0 1 0.8 0 0.0 0 0.0 1 0.4 3 2.1 0 0.0
Insect
Unidentified insect 104 18.7 43 14.3 13 11.0 48 35.3 1 1.8 59 24.5 16 11.2 28 23.7
Mammal
Odocoileus virginianus White-tailed Deer 35 6.3 16 5.3 15 12.7 4 2.9 5 8.9 11 4.6 7 4.9 15 12.7
Sus scrofa Feral Hog 9 1.6 5 1.7 2 1.7 2 1.5 0 0.0 4 1.7 2 1.4 3 2.5
Dasypus novemcinctus Armadillo 11 2.0 6 2.0 4 3.4 1 0.7 0 0.0 8 3.3 3 2.1 0 0.0
Procyon lotor Raccoon 162* 84* 42* 36* 11* 90* 45* 16*
Urocyon cinereoargenteus Gray Fox 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0
Sylvilagus floridanus EasternCottontail 47 8.5 21 7.0 5 4.2 21 15.4 2 3.6 24 10.0 9 6.3 12 10.2
Sylvilagus aquaticus Swamp Rabbit 2 0.4 1 0.3 1 0.8 0 0.0 0 0.0 0 0.0 2 1.4 0 0.0
Neotoma floridana Eastern Woodrat 4 0.7 1 0.3 3 2.5 0 0.0 1 1.8 2 0.8 1 0.7 0 0.0
Rattus norvegicus Norway Rat 1 0.2 0 0.0 1 0.8 0 0.0 1 1.8 0 0.0 0 0.0 0 0.0
Sigmodon hispidus Hispid Cotton Rat 32 5.8 22 7.3 6 5.1 4 2.9 1 1.8 18 7.5 10 7.0 3 2.5
Reithrodontomys fulvescens Fulvous Harvest Mouse 3 0.5 1 0.3 2 1.7 0 0.0 1 1.8 1 0.4 1 0.7 0 0.0
Ochrotomys nuttalli Golden Mouse 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0
Plant
Callicarpa americana Beauty Berry 24 4.3 23 7.7 0 0.0 1 0.7 13 23.2 0 0.0 9 6.3 2 1.7
Rubus plicatus Bramble Blackberry 100 18.0 57 19.0 21 17.8 22 16.2 1 1.8 77 32.0 22 15.4 0 0.0
Celtis spp. 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 0 0.0 0 0.0 1 0.8
Prunus virginiana Choke Cherry 12 2.2 5 1.7 7 5.9 0 0.0 6 10.7 1 0.4 5 3.5 0 0.0
Zea mays Corn 28 5.0 19 6.3 1 0.8 8 5.9 0 0.0 2 0.8 2 1.4 24 20.3
Unidnetified grasses 30 5.4 12 4.0 12 10.2 6 4.4 3 5.4 15 6.2 9 6.3 3 2.5
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Annual 2009 2010 2011 Fall Spring Summer Winter
Type/species Common name n % n % n % n % n % n % n % n %
Smilax rotundifolia Greenbriar 15 2.7 15 5.0 0 0.0 0 0.0 6 10.7 0 0.0 9 6.3 0 0.0
Celtis occidentalis Hackberry 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 0 0.0 0 0.0 1 0.8
Ilex spp. Holly 3 0.5 2 0.7 0 0.0 1 0.7 0 0.0 0 0.0 0 0.0 3 2.5
Morus nigra Mulberry 4 0.7 3 1.0 0 0.0 1 0.7 2 3.6 1 0.4 0 0.0 1 0.8
Vitis rotundifolia Muscadine 15 2.7 8 2.7 7 5.9 0 0.0 2 3.6 0 0.0 13 9.1 0 0.0
Ampelopsis arborea Peppervine 2 0.4 1 0.3 1 0.8 0 0.0 1 1.8 0 0.0 1 0.7 0 0.0
Diospyros virginiana Persimmon 10 1.8 6 2.0 4 3.4 0 0.0 6 10.7 0 0.0 4 2.8 0 0.0
Ilex decidua Possum Hawe 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 1 0.4 0 0.0 0 0.0
Cucurbita spp. Pumpkin 1 0.2 1 0.3 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 1 0.8
Lolium multiflorum Ryegrass 5 0.9 3 1.0 0 0.0 2 1.5 0 0.0 0 0.0 0 0.0 5 4.2
Citrullus lanatus Watermelon 2 0.4 0 0.0 2 1.7 0 0.0 0 0.0 0 0.0 2 1.4 0 0.0
Nyssa aquatica Water Tupelo 4 0.7 4 1.3 0 0.0 0 0.0 0 0.0 0 0.0 2 1.4 2 1.7
Unidentified plants 8 1.4 4 1.3 4 3.4 0 0.0 2 3.6 2 0.8 4 2.8 0 0.0
Invertebrates
Millipede 1 0.2 0 0.0 0 0.0 0 0.0 1 1.8 0 0.0 0 0.0 0 0.0
Cambarus spp. Crawfish 17 3.1 6 2.0 3 2.5 8 5.9 0 0.0 8 3.3 1 0.7 8 6.8
Aquatic snail 2 0.4 2 0.7 0 0.0 0 0.0 0 0.0 0 0.0 0 0.0 2 1.7
Micellaneous
Duck tape 1 0.2 0 0.0 0 0.0 0 0.0 0 0.0 1 0.4 0 0.0 0 0.0
Styrofoam 1 0.2 0 0.0 0 0.0 1 0.7 0 0.0 1